Secondary production of Calanus helgolandicus in the Western English Channel

The objective of this study was to investigate the production of Calanus helgolandicus off Plymouth, Western English Channel, from April to August 1998 comparing a range of different methods. The abundance of all developmental stages from NI to adult was determined. Body carbon and nitrogen and length were measured on individual NVI to adults. Female egg production was also monitored. All developmental stages were generally represented but four main periods of recruitment were determined. Two periods of low egg production were observed on the 13th and on the 27th of July: (i) the first was due to the arrival of newly moulted or not yet fertilised females in the population and (ii) the second low period was possibly due to the senescence of the females. Finally, the total production of C. helgolandicus was estimated using four different methods: the weight increment, the egg production, Huntley and Lopez and Hirst and Lampitt methods. Depending on the methods used, the production accumulated over the period from May 14 to August 10 ranged between 12 and 45 mg C m⁻³.     

The effect of food limitation on gonad development and egg production of the planktonic copepod Calanus finmarchicus

Calanus finmarchicus, one of the dominant copepods species of the North Atlantic, often encounters low food concentration or quality during the reproductive period; however, our knowledge on the effect of these conditions on reproduction processes is scarce. The present study combines experiments with histological observations to describe the response of C. finmarchicus to limited food focussing on (1) oocyte maturation processes, (2) gonad morphology, (3) egg production rates as a function of spawning frequency and clutch size, and (4) the fuelling of egg production by measuring carbon and nitrogen content of the females. In the laboratory, C. finmarchicus females were exposed to 0, 10, 50, 150 or >300 μg C l⁻¹ for several days. To account for food quality and season, reproductive activity was compared in April and July 1999 between females feeding on diatoms or dinoflagellates. The effect of feeding history was studied in February with females fed and starving prior to the experiment. Feeding conditions had severe effects on oocyte maturation process. Hence, egg production varied significantly with food concentration and quality, season and feeding history due to variation in both clutch size and spawning frequency. Clutch size differed by a factor of 2-4 between food limited and well fed females, and is thus an important parameter for modelling egg production. Changes in clutch size were related to changes of the number of maturing oocytes in the females gonads indicating that the latter can be used to precise the prediction of egg production from preserved samples. The proportion of females carrying at least some mature oocytes was relatively high at low food availability. Apparently, these females used internal body reserves as the carbon and nitrogen content decreased significantly under these conditions. These results indicate that C. finmarchicus embarks on the strategy to enable reproduction in all or many females of a population at low rates when feeding conditions are unfavourable.     

Effects of ultraviolet-B stressed diatom food on the reproductive output in Mediterranean Calanus helgolandicus (Crustacea; Copepoda)

Recent predictions anticipate a solar ultraviolet-B increase of 5-10% over temperate latitudes within the next 15 years. As Mediterranean waters are clear, enabling deep UVB penetration, we used controlled experiments to test the effects of equivalent levels of UVB on trophic plankton interactions. We grew the diatom Skeletonema costatum in the laboratory under daily 4 h UVB exposure in doses of 10% higher than today in contrast to control cultures (no UVB). UVB-exposed algae showed modification in cell structure, volume increase and delay in cell division, eventually leading to lysis and cell death after five days of daily 4 h exposure. Both control and UVB-stressed S. costatum cultures were used as food for wild Calanus helgolandicus females collected in the NW Mediterranean. Those fed on control diatoms produced three times more eggs, healthier offspring with fewer lethal naupliar deformities than females fed on UVB-exposed diatoms. Our results provide the first experimental evidence of indirect UVB effects on reproductive output through trophic response in marine plankton. The sensitivity of the S. costatum strain is discussed as well as the reduced reproductive output of the female copepods. We suggest that the low egg numbers and high naupliar deformity ratios are possibly a consequence of UVB-induced deficiency of essential components in the UVB-stressed maternal diet.     

Growth and development of Calanus chilensis nauplii reared under laboratory conditions: testing the effects of temperature and food resources

We assessed growth and development of naupliar stages of Calanus chilensis Brodsky 1959, under a combination of three temperatures and two food levels in laboratory conditions. Both food supply and temperature significantly affected naupliar growth and development. High food, measured as chlorophyll a (Chl a) concentration, was 40 μg l⁻¹, on average, and yielded temperature-dependent growth rates in the range of 0.13-0.17 day⁻¹. Low food was about 1.2 μg chlorophyll a l⁻¹ and retarded or arrested development and drastically reduced the growth rate to the range of 0.05-0.09 day⁻¹. To test whether these experimental results were consistent with field data, we used published information on temperature and chlorophyll a variability in northern Chile and developed a combined temperature/food-dependent model to diagnose naupliar growth in the field through a 2-year seasonal cycle including the 1997-1998 El Niño conditions. We concluded that in the upwelling region off northern Chile, C. chilensis might seldom encounter conditions of food shortage, as those applied in the laboratory. Thus, naupliar growth of this species may be primarily controlled by environmental temperature and this might also be the case for the dynamics of the entire population inhabiting the coastal upwelling zone.     

Effects of food nitrogen content and concentration on the forms of nitrogen excreted by the calanoid copepod, Acartia tonsa

Nitrogen excreted as ammonium, urea, and dissolved primary amines (DPA), and nitrogen ingested by the planktonic calanoid copepod, Acartia tonsa, were measured while fed 4 foods with different N/C ratios in high (500 μg C l^− 1) and low (50 μg C l^− 1) concentrations. Adult copepods were fed the ciliate, Uronema marinum (N/C = 0.26), the diatom, Thalassiosira weissflogii, in log-phase growth (N/C = 0.20), and in senescent-phase growth (N/C = 0.12), and detritus derived from the saltmarsh grass, Spartina alterniflora, (N/C = 0.04). Total nitrogen excreted ranged from 0.06 to 0.18 μg N copepod^− 1 d^− 1 whereas nitrogen ingested exhibited considerably more variation (0.01 to 0.39 μg N copepod ^− 1d ^− 1). Ammonium was the dominant form of nitrogen excreted and was influenced by both food concentration and N/C ratio. Copepods fed foods with N/C ratios resembling their own body composition (log-phase diatoms and ciliates) excreted more ammonium when fed higher concentrations of food. In contrast, copepods fed foods with lower N/C ratios than their own body composition excreted more ammonium when fed lower concentrations of food, suggesting that they were catabolizing body protein for survival. Excretion of urea varied with food N/C ratio, with more urea excreted when the copepods were fed higher N/C foods. The excretion of DPA did not vary with either food concentration or food N/C ratio. Homeostasis serves to conserve the N/C ratio of copepods. Thus nitrogen excretion by healthy copepods should be expected to increase with ingestion only when copepods have high quantities of nitrogen-rich foods relative to the body composition of the copepods.     

Effect of essential dietary fatty acids on egg production and hatching success of the marine copepod Temora longicornis

The calanoid copepod Temora longicornis and its food (seston of size < 200 μm) was sampled during three successive seasons (2002-04) in the Trondheimsfjord, Central Norway. Egg production (24 h) and hatching success (72 h) was determined by incubation experiments, and the essential fatty acid (EFA) content of their in situ food was analysed. The dominant EFA in the seston were DHA (22:6n-3) and EPA (20:5n-3), whereas ARA (20:4n-6) were present in low quantities. Egg production and hatching success was relatively low during early spring and late autumn (~ 10 eggs female^- 1 day^- 1 and ~ 30%), and relatively high during summer. Spring phytoplankton, dominated by diatoms, contained low amounts of DHA. Dinoflagellates, small flagellates, and ciliates dominated during summer, when a high content of DHA was recorded.The rate of egg production of T. longicornis did not show any relationship with food concentration (r² = 0.003), but was positively correlated to temperature, although not statistically significantly (r² = 0.48, p = 0.05). The quantitative and percent DHA contents of the food was significantly related to the rate of egg production (r² = 0.96 and 0.95, respectively, p < 0.001), but no such relationship were observed for the quantitative or percent content of EPA and ARA in the seston. The egg production of T. longicornis during May-August was 43-47 eggs female^- 1 day^- 1, with dietary DHA contents higher than 7-8 mg DHA g^- 1 DW. Also the hatching success of T. longicornis was positively correlated to the quantitative content of DHA in the diet (r² = 0.88), but hatching was also inversely related to the percentage ARA (r² = 0.84). The maximum hatching success was found when the ARA content was < 0.15% of total fatty acids and the DHA:ARA ratio was > 50. The conclusion that DHA most strongly affected egg production whereas ARA affected hatching, fit well with earlier findings for fish. Our results do not exclude that toxic aldehydes interact with reproduction of calanoid copepods when diatoms are present, but we observed a consistent pattern where dietary DHA and ARA alone explained a majority of the variability in egg production and hatching of T. longicornis.     

Application of a kinetic model to demonstrate how selective feeding could alter the amount of cadmium accumulated by the blue mussel (Mytilus trossolus)

Filter-feeding bivalves are well adapted to a changing food environment, capable of a highly selective feeding strategy. This feeding behaviour results in the rejection of unwanted material as pseudofeces. To assess the importance of a selective feeding behaviour performed by filter-feeding bivalves in influencing amounts of metal accumulated by bivalves, we applied a kinetic model of metal accumulation modified to incorporate feeding behaviour defined as (1) the sorting efficiency (SEF; the ability of the mussel to select organic matter (OM) over inorganic matter (IM) from suspended particulate matter (SPM) cleared from the water column) and (2) rate of rejection of unwanted SPM as pseudofeces (PF). Predicted mussel Cd concentrations (Cd_muss) were compared with Cd_muss predicted without incorporating feeding behaviour as well as to observed Cd_muss. SEF was important at an SPM quality greater than 10% OM; PF production was important at an SPM quantity greater than 3.5 mg l⁻¹ with the influence of PF production on predicted Cd_muss increasing with increasing SPM concentration. Our results suggest that a selective feeding behaviour and the generation of pseudofeces could influence amounts of metal ingested, especially under conditions where SPM quality and quantity are highly variable. Under conditions of high SPM, the influence of SEF and PF production on final Cd_muss should be taken into consideration.     

Toxin content and toxic effects of the dinoflagellate Gyrodinium corsicum (Paulmier) on the ingestion and survival rates of the copepods Acartia grani and Euterpina acutifrons

Short-term experiments showed that ingestion rates of the copepods Acartia grani and Euterpina acutifrons on different concentrations of the dinoflagellate Gyrodinium corsicum were not usually different from those obtained with the non-toxic and similar sized Prorocentrum triestinum. Long-term experiments showed that no A. grani survived for more than 288 h on concentrated cultures (1500 μg C l⁻¹) of G. corsicum compared with survival rates of 86.7% on the non-toxic P. triestinum. In contrast high and similar survival rates were found for E. acutifrons exposed for the same time to similar concentrations of both dinoflagellates. These results demonstrated that G. corsicum produce toxins which have significant adverse effects on the long-term survival rates of some copepods like A. grani but not on other copepods like E. acutifrons. The possibility of PSP-toxin production by G. corsicum was analysed by chromatographic analysis (HPLC-FD) and mouse bioassay of extracts obtained from cultures maintained in exponential growth phase in f/2-Si and L1 culture mediums. These analyses and the bioassay ruled out the presence of PSP toxins in this dinoflagellate. However, mouse tests provided the first evidence for the presence of some type of NSP toxin in G. corsicum. Haemolytic assays also gave the first positive results for the methanol extract of this dinoflagellate species.     

Nucleic acid indices of egg production in the tropical copepod Acartia sinjiensis

Two experiments were conducted to evaluate the effects of food and temperature on the nucleic acid content and egg production (and their relationship) of the tropical copepod Acartia sinjiensis. Experiment 1 evaluated the effect of food quality (as different algae species) on the relationship between nucleic acid content and egg production. In Experiment 2, the main and interaction effects of food type, food concentration and temperature on the total, Carbon and Nitrogen specific egg production and nucleic acid indices were evaluated in a factorial experimental design. Food quality, concentration and temperature significantly affected the nucleic acid content, egg production rate and the nucleic acid-egg production relationship of A. sinjiensis. RNA indices were correlated with egg production in females fed Pavlova salina, Tetraselmis chuii and Chaetoceros muelleri, but not in females fed Isochrysis aff. galbana. The slopes of the linear regressions of RNA indices as predictors of egg production were similar in females fed different algae species, suggesting that the slope of these relationships might be independent of food quality. The DNA content of females was significantly affected by food and temperature, suggesting that it is not a good index of cell number in this species. Nevertheless, the RNA:DNA ratio was as good a predictor of egg production as total RNA content. Egg production showed a weakly positive correlation with temperature. On the other hand, total, C- and N-specific nucleic acid indices had a strong negative correlation with temperature. In addition, temperature had a non linear effect on the slopes of the regression lines of RNA content and RNA:DNA ratio as predictors of egg production—slopes were similar at 25 °C and 30 °C, but significantly lower at 20 °C. Furthermore, the predictability of egg production was improved when the interaction term of nucleic acid indices with temperature was used instead of the nucleic acid indices alone in linear regression models. Our results suggest food quality has a limited influence on the nucleic acid-egg production relationship, and that temperature should be accounted for in models using nucleic acid indices as predictors of egg production in A. sinjiensis.     

Effects of estuarine conditions and organic enrichment on the fecundity and hatching success of Acartia clausi in contrasting systems

The fecundity and hatching success of Acartia clausi were analysed at fixed salinity sites (35, 34 and 33 psu) in two nearby estuaries (Bilbao and Urdaibai, Basque coast, Bay of Biscay) from March to June 1997. Field incubations were conducted to estimate egg production rates and hatching success, and the size of eggs and experimental females measured. Water temperature and dissolved oxygen saturation were also determined, as well as seston samples to quantify food abundance and quality. Between-estuary and within-estuary differences were tested statistically, and correlation and regression analyses were used to determine relationships between reproductive and environmental variables. Egg production rates were higher in the organically enriched estuary of Bilbao; this denoting that food supply controls the fertility of A. clausi in these systems. Temporal patterns of egg production differed between estuaries, and were associated with different nutritional factors in each estuary. Within the salinity range analysed, egg production reached higher values at intermediate salinity (≈34 psu) in both estuaries. This was interpreted as the result of the interaction between the positive effect of food increase, and the negative effect of physicochemical conditions with decreasing salinity. Egg size variations mainly occurred temporally in relation to female size, but no clear trade-off between egg size and egg number was observed in any case. A drop in hatching success in Bilbao, mainly in waters of <34 psu, was related to the oxygen depletion caused by organic pollution. This indicates that organic enrichment in Bilbao has opposite effects on the reproductive success, because it enhances egg production but reduces offspring survival.     

Intraspecific egg size variation and sperm limitation in the broadcast spawning bivalve Macoma balthica

Bottom-ice algae within Antarctic sea ice were examined using chlorophyll fluorescence imaging. The detailed structure of the bottom-ice algal community growing in the platelet and congelation layers of solid pieces of sea ice was evident for the first time in chlorophyll imaging mode. Strands of fluorescence representing algal cells were clearly visible growing upward into brine channels in a fine network. Images of effective quantum yield (Ф_PSII) revealed that the Ф_PSII of algae embedded in the sea ice was approximately 0.5. Furthermore, Ф_PSII decreased slightly with distance from the ice-water interface.The response of Antarctic sea ice algae to changes in irradiance and salinity, and the effects of slowly warming and melting the ice block sample were examined using this system. The Ф_PSII of bottom-ice algae decreased as irradiance increased and salinities decreased. Bottom-ice algae appear to be most vulnerable to changes in their environment during the melting process of the ice, and this suggests that algae from this region of the ice may not be able to cope with the stress of melting during summer.Chlorophyll fluorescence imaging provides unprecedented imagery of chlorophyll distribution in sea ice and allows measurement of the responses of sea ice algae to environmental stresses with minimal disruption to their physical habitat. The results obtained with this method are comparable to those obtained with algae that have been melted into liquid culture and this indicates that previous melting protocols reveal meaningful data. In this chlorophyll imaging study, rapid light curves did not saturate and this may prevent further use of this configuration.     

Mating and role of seminal receptacles in the reproductive biology of the spider crab Maja squinado (Decapoda, Majidae)

The reproductive biology of the spider crab Maja squinado was analyzed based on monthly samples from an 18-month study carried out in Galicia (NW Spain) and laboratory experiments holding primiparous and multiparous females in captivity with and without males. The seminal receptacles of adult females were analyzed and their relationship with the presence and developmental stage of the eggs and the gonad maturity stage was determined. Gonad maturation in primiparous females began one or two months after the pubertal moult. Females having gonads in an advanced stage of development made their appearance in December and the first spawning took place in mid-winter or early spring. The percentage of ovigerous females from March to September was ∼75%. As the incubation period progressed, the ovaries became mature again in order to carry out the next spawning. Under experimental conditions the breeding cycle started earlier in multiparous females, during their second yearly cycle, than in primiparous ones. After mating, female spider crabs store sperm in seminal receptacles and this sperm is used in the fertilization of eggs immediately prior to spawning. The analyses of seminal receptacles consisted of the estimation of fullness and the number of differentiated sperm masses. The number of masses ranged between 0 and 6 in field samples (median for females with stored sperm=1) and was positively correlated with fullness. Differences in colour and volume of individual masses showed that, at least in some cases, females carried out successive matings with long intervals in between. This storage mechanism allowed females to fertilize successive broods without remating (as was also shown under experimental conditions). Juvenile females from shallow waters did not have developed seminal receptacles which indicated that mating was not possible until the onset of maturity. Postpubertal females in shallow waters (August to October), including animals participating in aggregations, always showed empty receptacles. The seasonality of receptacle fullness showed that mating involved hard-shelled females and occurred in deep water during the autumn migration from juvenile habitats or in the wintering habitats, during the last stages of gonad maturation (November to February). After fertilization ovigerous females continued to store sperm, but the volume was lower than in non-ovigerous females. Mating may occur in ovigerous females, particularly in the final period of incubation, because in females with broods almost ready to hatch, both new and older sperm masses were seen in the receptacles (distinguished by colour and size). The fullness of the receptacles decreased both in ovigerous and non-ovigerous females in the final phase of the annual breeding cycle (August–October), however, some sperm was still available. In the laboratory, mating was observed, and no courtship nor postcopulatory guarding was recorded. The analysis of receptacles from laboratory experiments indicated that primiparous and multiparous females showed differences in the seasonality of mating in the first phase of the breeding cycle (September–January), related to differences in the timing of gonad maturation and hatching. Mating occurred in the final stages of gonad maturation, a short time before hatching, and matings were detected in ovigerous females. Multiple matings were also evident, to a greater extent than in the field, probably due to the higher availability of males. Females underwent over four successive spawnings in the laboratory without having to recopulate, and the incubation lasted on the average from 40 to 58 days (∼18 and 16°C respectively) and the mean duration between hatching and the next spawning was 3.4 days. It is estimated that most females carry out three successive spawnings during the annual cycle.     

The influence of an intermittent food supply on the feeding behaviour of Yoldia hyperborea (Bivalvia: Nuculanidae)

Yoldia hyperborea (Lovén) is a protobranch bivalve of circumboreal distribution and an important bioturbator of muddy sediments of cold-water embayments in Newfoundland, Canada, where it is exposed to a strong seasonal input of sinking phytoplankton during spring, sporadic events of sediment resuspension, and sediment of low nutritional value during winter. To explain field-observed patterns of population dynamics, we quantified the behavioural response of Y. hyperborea to pulses of settling algae and sediment resuspension events, and hypothesised that Y. hyperborea behaviour is modified during settling of nutrient-rich organic matter. Yoldia hyperborea responded rapidly to the arrival of settling microalgae by extending its siphons into the water column. Once the pulse of algal material reached the sediment the animals partially emerged, extended the palp proboscides over the sediment surface and maintained close contact with the area of highest algal concentration. In contrast, the activity of animals not exposed to settling algae or to resuspended sediment was primarily restricted to strata below the sediment surface. Thus the predominant subsurface feeding behaviour of Yoldia hyperborea is switched to surface deposit-feeding as a response to cues contained in microalgae. Results suggest active suspension-feeding during algal sedimentation events, although deposit-feeding resumes once the algae are no longer in suspension, and suspension-feeding is probably of little nutritional significance. The rapid behavioural response of Y. hyperborea to the influx of high quality food, such as fresh microalgae, is probably an adaptive foraging strategy to a food-limited environment and suggests higher bioturbation rates of surface sediments during spring, a key role being played by the protobranch in redistribution of labile phytodetritus within the benthos.     

Different diatoms induce contrasting effects on the reproductive success of the copepod Temora stylifera

Four diatoms, Thalassiosira rotula (THA), Thalassiosira weissflogii (TWEI), Skeletonema costatum (SKE) and Phaeodactylum tricornutum (PHA), and one dinoflagellate Prorocentrum minimum (PRO, control diet), were tested during the course of 14-day laboratory experiments examining the effects of these algae on the fecundity and egg viability in the copepod Temora stylifera. All algae were provided at the same mean carbon concentrations of 0.98 μg C ml⁻¹ to normalize the effects due to differences in diatom size and nutrient concentrations, at least with regards to carbon, so that variations in egg production and hatching viability were presumably due to other chemical constituents of the cells. Our results show that only PRO supported high reproductive success, with stable egg production rates and egg viability close to 100%. By contrast, all four diatoms negatively impacted egg production and hatching success, with varying degrees of suppression. The two diets SKE and PHA induced dramatic effects on reproductive success; SKE was the worst diet with naupliar production ceasing altogether within only 3 days. With PHA, fecundity never reached zero, but at the end of the experiments, initial egg production rates had dropped to 6.6% of initial rates, and viability had declined to zero within 7 days. The two Thalassiosira species mainly affected egg viability, so that after 14 days of feeding, a decrease of 90% had occurred. However, egg production rates were high and were reduced only by 15-24% towards the end of the experiments. In terms of potential recruitment rates, even though PRO induced high mean naupliar production, a slight decrease was observed during the first 5 days, likely due to the time required for T. stylifera to acclimate to PRO. After day 5, potential recruitment rate was higher, comparable to in situ values. THA and TWEI inhibited potential recruitment up to 90-100%, but these effects were not evident over a short-term period, denoting the importance of introducing the time factor when evaluating diatom effects on copepod production. With the other two diatom diets, SKE and PHA, recruitment was zero after 2 and 6 days, respectively. Experiments testing the effects of unsaturated aldehydes in THA (2-trans-4-trans-decadienal) and SKE (2-trans-4-trans-octadienal and 2-trans-4-trans-heptadienal) on the hatching viability of T. stylifera indicate that diatom-derived aldehydes may differ in terms of biological activity. Of the three molecules tested, decadienal had somewhat stronger effects on hatching success, with total blockage occurring at concentrations of 2.0 μg ml⁻¹; the concentrations to induce total blockage by octadienal and heptadienal were somewhat higher and ≥2.5 μg ml⁻¹. We suggest that the greater biological activity of SKE in terms of cell blockage may thus be due to the presence of antimitotic compounds other than these unsaturated aldehydes.     

Egg production of the copepod Acartia bifilosa in two contrasting European estuaries in relation to seston composition

The egg production of the copepod Acartia bifilosa was measured and related to environmental variables and food availability in two estuaries located in the same biogeographic region (Bay of Biscay) but showing very strong differences in abiotic and biotic features: the Gironde estuary (France) and the estuary of Mundaka (Spain). The study was conducted during the spring-summer-autumn period of 1994. Food availability was evaluated by analysing the chlorophyll a (Chl a), the particulate organic carbon (POC) and the easily extractable macromolecular compounds such as proteins, carbohydrates and lipids of the seston. The egg production of copepods was estimated from field incubations with natural water, and phytoplankton feeding of adult females was estimated by means of the gut fluorescence method. The nutritional environment of the Gironde was characterised by high amounts of suspended particulate matter (SPM) with low food value, emphasising the mainly detrital origin of the organic matter (OM). In Mundaka, the higher contribution of phytoplankton to the seston led to marked increases in particulate food value accounting for up to 35% of organic matter. The weight-specific egg production was found to be sharply higher in Mundaka (ranging from 0.2 to 0.63×10⁻³ day⁻¹) than in the Gironde (ranging from 0 to 0.13×10⁻³ day⁻¹), but the seasonal trend of variations was similar, the highest weight-specific egg production rates occurring in early summer and the lowest in autumn in both estuaries. Egg production was not correlated linearly with temperature since maximal egg production occurred at intermediate temperatures. In Mundaka, the egg production showed a significant positive correlation with the chlorophyll and the Chl/SPM and the POC/SPM ratios. This coupled with higher values of algal food availability (Chl a/SPM: 10 to 1870 μg g⁻¹) and gut fluorescence (between 0.12 and 0.38 ng Chl a Eq ind⁻¹) indicate that a herbivorous diet could cover the energy requirements of A. bifilosa and support egg production. In the Gironde, the algal food availability and the gut fluorescence were lower (Chl a/SPM: 10 to 80 μg g⁻¹; GF: 0.09 and 0.25 ng Chl a Eq ind⁻¹), and the egg production showed significant positive correlation with the particulate food value, suggesting that other sources of carbon rather than phytoplankton were responsible for the observed changes in egg production. Results indicate that the particular seston properties of each system may be responsible for the noticeable differences in A. bifilosa fertility among estuaries.     

Effects of different diets on the reproduction and naupliar development of the copepod Acartia bifilosa

The influence of diatoms on the reproduction and naupliar development of Acartia bifilosa was investigated under laboratory conditions, comparing initial in situ values and laboratory-food treatments. Egg production by A. bifilosa was significantly reduced by one diatom diet (Phaeodactylum tricornutum: Pt) and by two non-diatom diets (Platymonas subordiformis: Ps and Nannochloropsis oculata: No). It was less affected by the other diatom diet (Skeletonema costatum: Sc) or by two mixed-food treatments (D-mix and DG-mix), composed of two diatoms (Pt, Sc) and four species (Pt, Sc, Ps and No), respectively. The negative effect of Pt was eliminated when adult copepods were offered mixed-food diets. There were no significant differences between the hatching success values observed in filtered seawater and in algal exudates, indicating that diatoms did not produce active dissolved toxic substances under the different food concentrations tested. The mortality rate of nauplii was higher with Pt than the other diets, suggesting that this diatom species had a negative effect on egg production, hatching success and naupliar survival simultaneously. Compared to other diets, No and Pt were not beneficial food sources for reproduction and for female and larval survival.     

Comparison of diet and feeding activity of two polychaetes, Nephtys caeca (Fabricius) and Nereis virens (Sars), in an estuarine intertidal environment in Québec, Canada

It is crucial to study how food resources are allocated in order to understand how invertebrate species of soft-bottomed benthic communities can live in the same environment, especially when belonging to the same trophic level. On the southern shore of the Lower St. Lawrence Estuary, the polychaetes Nereis virens (Nereidae) and Nephtys caeca (Nephtyidae) represent the main infaunal predators and are known to regulate (by predation) other populations within their community. For these animal populations, there is a high degree of spatial overlap in the horizontal dimension as well as vertically in the sediment column. The purpose of this study was to describe and compare the feeding activity of N. virens and N. caeca in relation to several environmental factors. The feeding activity of N. caeca was further studied under natural and experimental conditions and a feeding experiment was carried out to evaluate how the food resource is subdivided in conditions of high spatial overlap. Finally, the prospecting activity of N. virens was described under natural conditions. N. virens and N. caeca exhibited a high degree of dietary overlap, particularly at the adult stage. This food resource overlap can easily influence the regulation of these species, even though they display different feeding responses to environmental stimuli. Tides and storm events can also influence the feeding behaviour. We conclude that different behavioural feeding responses likely favour the occurrence of these two species in the same trophic environment.     

铜绿微囊藻和斜生栅藻非稳态营养盐限制条件下的生长竞争特性

采用“脉冲”添加方法进行了非稳态条件下铜绿微囊藻（M.）和斜生栅藻（S.）分别在氮磷单营养盐和双营养盐限制时的共培养试验。试验结果显示：当添加频率为1d时,无论何种营养盐限制,M.均成为优势藻种。氮限制条件下,氮时均浓度范围在0.3—2.4 mg/L时,M.始终具有竞争优势。磷限制条件下,磷浓度范围在0.018—0.035 mg/L时,S.只在生长初期阶段占优。氮磷双营养盐限制条件下,添加液的氮磷质量比为35:1（设定为最优比）,添加频率为8 d时,两种藻表现出共生特征;而偏离最优比时（N:P=70:1,17:1）,在不同的添加频率下均未出现共生现象,且氮的时均浓度为0.6—4.8 mg/L时（70:1）,M.具有竞争优势,而降低为0.15—0.3 mg/L时（17:1）,S.占优。随着添加频率的变化,两种藻的细胞大小也会随之改变,S.随着营养盐浓度的降低而增大,且在双营养盐限制条件下变化更显著。上述试验结果分析表明：两种藻竞争能力与添加频率相关,在藻种浓度的变化上,按照‘拾遗－机会’交替竞争理论,M.表现出机会主义者特征,而S.则表现出拾遗者的特征,两者的共生特征也符合‘中度干扰’假说。藻细胞大小变化表明,两种藻均可以改变大小实现最大限度争夺受限营养盐。在低浓度时,S.细胞大小的变化同样也变现出了“拾遗者”的特征。     

脂肪酸对中华哲水蚤摄食两种海洋微藻的指示作用

在室内以饥饿培养为对照,以海洋原甲藻(Prorocentrum micans)和中肋骨条藻(Skeletonema costatum)培养中华哲水蚤(Calanus sinicus),研究了各脂肪酸标记对中华哲水蚤摄食不同饵料的指示作用。结果显示,海洋原甲藻中18 ∶ 4ω3、22 ∶ 6ω3含量较高,中肋骨条藻中16 ∶ 1ω7、20 ∶ 5ω3的含量较高。二者分别表现出典型的甲藻门和硅藻门的脂肪酸组成特征。中华哲水蚤的脂肪酸组成有两个特点:(1)20 ∶ 5ω3和22 ∶ 6ω3的含量均较高;(2)其体内表征桡足类浮游植物食性、由桡足类自身合成的20 ∶ 1和22 ∶ 1脂肪酸占有相当的比例。虽然中华哲水蚤对不同脂肪酸的吸收和转化效率不同,但以脂肪酸作为标记还是成功的指示了中华哲水蚤对微藻的摄食。在饥饿培养中,首先消耗的是那些浮游动物自身不能合成的多不饱和脂肪酸,而结构脂肪酸都表现出了较高的保守性。结合各脂肪酸标记变化趋势和Pearson相关性分析的结果认为,18 ∶ 4ω3、18 ∶ 4ω3/16 ∶ 1ω7、∑18/∑16能较好的指示中华哲水蚤对海洋原甲藻的摄食,仅16 ∶ 1ω7/18 ∶ 4ω3能指示中华哲水蚤对中肋骨条藻的摄食。     

Responses of the black sea urchin Tetrapygus niger to its sea-star predators Heliaster helianthus and Meyenaster gelatinosus under field conditions

We ran field experiments to examine the responses of the black sea urchin Tetrapygus niger to predatory sea stars. Trials involving simulated attacks (one or several arms of a sea star being placed on top of half the urchin) showed that the urchin differentiated between the predatory sea stars, Heliaster helianthus and Meyenaster gelatinosus, and a non-predatory sea star, Stichaster striatus, and showed almost no response to a sea star mimic. We further compared the responses of the urchin to different threat levels presented by the two predatory sea stars. The highest threat level was a simulated attack, then mere contact, and subsequently sea stars being placed at different distances from the urchin. All urchins responded to simulated attacks and contact with both sea stars. The proportion responding decreased with distance and more rapidly in trials with H. helianthus (0% at a distance of 30 cm) than with M. gelatinosus (33% at a distance of 50 cm). At each of the threat levels where there was a response to both sea stars, the urchins responded more rapidly to M. gelatinosus than to H. helianthus. In a third experiment where a predatory sea star was added to a circular area (1-m diameter) in which either 4-8 or 11-19 undisturbed urchins were present, the urchins fled the area more rapidly when the added sea star was M. gelatinosus, but the rate of fleeing did not vary with density, as might occur if there was communication among urchins using alarm signals. Our observations suggest that M. gelatinosus presents a stronger predatory threat than H. helianthus. This corresponds to field observations showing that the urchins are more frequently consumed by M. gelatinosus. These are the first field experiments demonstrating distance chemodetection by a marine invertebrate under back-and-forth water flow from wave activity.