The diatom Thalassiosira rotula affects reproductive success in the copepod Acartia clausi

Fecundity, egg viability and fecal pellet production are reported for Acartia clausi females collected in the Bay of Naples, Italy, from April to October 1992 and fed either with a diatom (Thalassiosira rotula) or dinoflagellate (Prorocentrum minimum) diet, at food saturated conditions. The diatom diet significantly reduced both egg and fecal pellet production as well as hatching success. Blockage of egg development occurred with both axenic and non-axenic cultures of T. rotula, suggesting that inhibitors were provided by the diatoms and not by the bacteria associated with diatom cultures. Low hatching success was also artificially induced by exposing newly spawned A. clausi eggs to high concentrations of diatom extracts, indicating the presence of deleterious, inhibitory compounds blocking copepod embryogenesis. Fecundity and hatching success diminished significantly with female age. In contrast, female longevity was not significantly modified by food type. The presence of males did not significantly alter fecundity or egg viability. Females continued to produce viable eggs throughout the period of incubation, with and without males, in both food conditions, indicating that remating is infrequent and not necessary to sustain viable egg production in this species. The succession in low and high population densities may therefore be the outcome of variations in survival rates of eggs, rather than reproductive protential perse; such variations may strongly depend on the adult copepod diet.     

Copepod recruitment and food composition: do diatoms affect hatching success?

Laboratory experiments were conducted to differentiate between factors controlling the hatching success of copepod eggs. Factors that could affect viability of eggs; viz food quality, female condition and external factors were investigated. In a series of experiments the copepod Acartia tonsa Dana was fed several different diets while egg production and hatching success were monitored. The diet was analysed for fatty acid content as an indicator of food quality. Both egg production and hatching were found to be affected by the nutritional quality of the food. Hatching was also highly dependent on female fertility. External effects were tested by exposing eggs to diatom extracts. Negative effects were only evident at high extract concentrations, but disappeared when aeration was supplied to the solution. Oxygen measurements showed that failure to hatch was due to hypoxia in the extracts. No inhibitory or toxic effects of diatom cell components on hatching could be found.     

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.     

Strain-related physiological and behavioral effects of <Emphasis Type="Italic">Skeletonema marinoi</Emphasis> on three common planktonic copepods

Three strains of the chain-forming diatom Skeletonema marinoi, differing in their production of polyunsaturated aldehydes (PUA) and nutritional food components, were used in experiments on feeding, egg production, hatching success, pellet production, and behavior of three common planktonic copepods: Acartia tonsa, Pseudocalanus elongatus, and Temora longicornis. The three different diatom strains (9B, 1G, and 7J) induced widely different effects on Acartia tonsa physiology, and the 9B strain induced different effects for the three copepods. In contrast, different strains induced no or small alterations in the distribution, swimming behavior, and turning frequency of the copepods. 22:6(n-3) fatty acid (DHA) and sterol content of the diet typically showed a positive effect on either egg production (A. tonsa) or hatching success (P. elongatus), while other measured compounds (PUA, other long-chain polyunsaturated fatty acids) of the algae had no obvious effects. Our results demonstrate that differences between strains of a given diatom species can generate effects on copepod physiology, which are as large as those induced by different algae species or groups. This emphasizes the need to identify the specific characteristics of local diatoms together with the interacting effects of different mineral, biochemical, and toxic compounds and their potential implications on different copepod species.     

Reproductive biology of the planktonic copepod Temora stylifera

Laboratory studies on the reproductive cycle of the planktonic copepod Temora stylifera, collected from the Gulf of Naples between October 1987 and March 1988, showed that females oscillated between a light and dark gonadal condition. Histological preparations indicated that the dark condition corresponded to a ripe stage of oogenesis as compared to the light or unripe maturation state of primary oocytes. The number of eggs released within 24 h was strongly dependent on the reproductive status of the female at the time of incubation. Light unfed females rarely reverted to a dark or ripe condition and egg deposition in such cases was almost always zero. Dark females produced eggs within 24 h under all experimental food conditions, reverting from dark to light soon after egg deposition. Successively, the length of the interclutch period was strongly dependent on the type of food present. A high percentage of infertile eggs was almost always present. We suggest that short-term studies, not taking into consideration the condition of gonads at the time of incubation, will underestimate potential egg production rates. Females maintained for longer than three or four days produced mainly infertile eggs. After remating, females reinitiated the production of fertile eggs but egg viabilities never reached 100% hatching success, indicating poor efficiency in the fertilization of mature oocytes. We hypothesize a possible cause and effect relationship between egg viabilities and seasonal fluctuations in population densities.Died tragically at sea during a cruise     

Extensive cross-disciplinary analysis of biological and chemical control of Calanus finmarchicus reproduction during an aldehyde forming diatom bloom in mesocosms

Egg and faecal pellet production and egg hatching success of the calanoid copepod Calanus finmarchicus were monitored over a period of 14?days (14?C28 April, 2008) while fed water from 4 differently treated mesocosms and ambient water. Two of the mesocosms used were inoculated with the polyunsaturated aldehyde (PUA)-producing diatom Skeletonema marinoi, while 2 received only nutrient additions with or without silica. The mesocosms developed blooms of S. marinoi, mixed diatoms or the haptophyte Phaeocystis pouchetii, respectively. Faecal pellet production of C. finmarchicus increased with increasing food availability. Egg production increased with time in all mesocosms to a maximum single female production of 232 eggs female^?1?day^?1 (average of 90 eggs female^?1?day^?1) and followed the development of ciliates and P. pouchetii, but was not affected by the observed high (up to 15?nmol?L^?1) PUA production potential of the phytoplankton. The hatching success of the eggs produced on the mesocosm diets was high (78?C96%) and was not affected by either aldehydes in the maternal diet or exposure to the dissolved aldehydes in the water.     

Phytoplankton food quality determines time windows for successful zooplankton reproductive pulses

Recruitment success at the early life stages is a critical process for zooplankton demography. Copepods often dominate the zooplankton in marine coastal zones and are prey of the majority of fish larvae. Hypotheses interpreting variations of copepod recruitment are based on the concepts of "naupliar predation," "nutritional deficiency," and "toxic effect" of diatom diets. Contradictory laboratory and field studies have reached opposite conclusions on the effects of diatoms on copepod reproductive success, blurring our view of marine food-web energy flow from diatoms to higher consumers by means of copepods. Here we report estimates of copepod feeding selectivity and reproduction in response to seasonally changing phytoplankton characteristics measured in a highly productive coastal upwelling area off the coast of central Chile. The variable phytoplankton diversity and changing food quality had a strong and highly significant impact on the feeding selectivity, reproduction, and larval survival of three indigenous copepod species. Seasonal changes in copepod feeding behavior were related to the alternating protozoan-diatom diets, mostly based on dinoflagellates and ciliates during winter and autumn (low highly unsaturated fatty acids [HUFA]/polyunsaturated fatty acids [PUFA] availability), but switched to a diet of centric and chain-forming diatoms (high HUFA/PUFA availability) during the spring/summer upwelling period. Ingestion of diatom cells induced a positive effect on egg production. However, a negative relationship was found between egg hatching success, naupliar survival, and diatom ingestion. Depending on the phytoplankton species, diets had different effects on copepod reproduction and recruitment. In consequence, it seems that the classical marine food web model does not apply to some coastal upwelling systems.     

On the life history of Centropages typicus: Responses to a fall diatom bloom in the New York Bight

Laboratory studies and field collections show that egg production by Centropages typicus (Krøyer) in New York shelf waters in autumn 1984 responded to both food and temperature. Rates of egg production were high (43 to 76 eggs female^-1 d^-1) in October, early in the fall diatom bloom. Later, although food concentrations remained high and female size actually increased, egg production declined, presumably in response to seasonally decreasing temperatures. Carnivorous diets did not support egg production. Development time for autumn-hatched C. typicus was 33 d at 15°C, a rate that gives a Q₁₀ of 2.21 when compared with the spring development rate of 49 d at 10°C. We could find no evidence of physiological adjustments being made by this copepod for overwintering. Development was not arrested at any subadult stage and resting eggs were not produced. Trends in body size of copepodid stage V, however, suggest that an overwintering strategy may be invoked by this copepod in Junuary or February.     

Fate of domoic acid ingested by the copepod <Emphasis Type="Italic">Acartia clausi</Emphasis>

Two important issues in the studies of harmful algae include ecological role of the toxic compounds and their fate through the food web. The aims of this study were to determine whether the production of domoic acid is a strategy evolved to avoid predation and the role of copepods in the fate of this toxic compound through the food web. The copepod Acartia clausi was fed with single and mixed cultures of the toxic diatom Pseudo-nitzschia multiseries and the non-toxic diatom Pseudo-nitzschia delicatissima. Ingestion rate as a function of diatom abundance was the same for the toxic and non-toxic Pseudo-nitzschia species, indicating no selective feeding behaviour against P. multiseries. The toxins ingested by the copepods did not affect mortality, feeding behaviour, egg production and egg hatching of the copepods. Copepods assimilated the 4.8% of the total domoic acid ingested. Although the amount of toxins daily detoxificated by the copepods was 63.6%, the copepods accumulated domoic acid in their tissues. We conclude that domoic acid is not toxic for copepods and, probably for this reason, this toxin does not act as feeding deterrent for copepods. However, even though the production of domoic acid has apparently not evolved to deter predation, copepods may play an important role on the fate of this toxic compound through the marine food web.     

Overwintering strategies of dominant calanoid copepods in the German Bight, southern North Sea

Abundance, stage composition and reproductive parameters (egg production, egg viability, proportion of spawning females) of the four copepod species Acartia clausi, Centropages hamatus, C. typicus and Temora longicornis were measured at the long term sampling station Helgoland Roads (German Bight, southern North Sea) from September 2003 to May 2004 to study their overwintering strategies. A. clausi was overwintering as females with arrested reproduction from November to January. T. longicornis, which is known to produce resting eggs in the North Sea, had a pelagic population with all developmental stages present during winter and reproductive rates closely related to food concentrations. Although their females produced eggs in response to ambient food conditions, both C. hamatus and C. typicus were rare in the pelagic. The C. hamatus population returned in May, probably from resting eggs, whereas C. typicus depended on advection. The Centropages species seemed to be less adapted to pelagic life in winter than A. clausi and T. longicornis. Sporadic occurrence of large numbers of nauplii and young copepodids of A. clausi and Centropages spp. pointed to different overwintering strategies or more successful survival in adjacent regions and advection of them into the waters around Helgoland island. While A. clausi was decoupled from environmental conditions in late autumn and winter, the other species were able to respond to variations in the food environment. Thus, egg production of T. longicornis increased during an unusual autumn diatom bloom.     

Are monoalgal diets inferior to plurialgal diets to maximize cultivation of the calanoid copepod Temora stylifera?

Temora stylifera adult copepods were fed with four different monoalgal diets and six combinations of the same cultures for 15 days. Fecundity, hatching success, number of cannibalized embryos, fecal pellet production, adult mortality and naupliar recruitment were compared, in order to find the best diet for this species. Phytoplankton species tested were Prorocentrum minimum (PRO); Isochrysis galbana (ISO); Tetraselmis suecica (TETRA) and Rhodomonas baltica (RHO) which were supplied alone or in different combinations and at various concentrations ranging from a minimum of 1 mg C L⁻¹ day⁻¹ to a maximum of 66 mg C L⁻¹ day⁻¹. Of the ten diets tested, ISO was the worst and was unable to sustain egg production and adult survival possibly because adults were unable to ingest this alga due to its small size. TETRA was also a poor food since it negatively impacted egg production and adult survival, as well as egg hatching success, possibly due to the lack of essential compounds necessary for optimal embryogenesis. RHO and PRO were the best foods inducing highest egg production, hatching success and naupliar recruitment. Even if mean egg production rates were similar to those obtained with some mixed diets, carbon intake concentrations with mixed diets were from 3 to 33 and from 6.6 to 66 times higher than with RHO and PRO given alone, respectively. Mixed diets of ISO and PRO, especially when supplied at higher concentrations (66 mg C L⁻¹ day⁻¹), had a negative effect on egg hatching success and adult survival, with a corresponding reduction in naupliar recruitment. On the other hand, mixed diets of TETRA and PRO promoted high naupliar recruitment but values were similar to PRO offered alone. Our results indicate that a good monoalgal diet such as RHO and PRO can be as effective as a mixed diet to sustain the mass cultivation of T. stylifera.     

Effects of season and maternal food on<Emphasis Type="Italic"> Calanus finmarchicus</Emphasis> reproduction,with emphasis on free amino acids

Effects of maternal food environment and season were examined during spring and autumn on females, eggs and nauplii of Calanus finmarchicus, in different natural prey suspensions or cultures of Rhodomonas baltica. Females sampled in spring were in general larger, had higher protein content, and showed higher egg production and hatching rates, than in autumn. The cumulative egg production was almost double in spring compared to autumn (females fed R. baltica). Females had higher content of free amino acids (FAA) and free essential amino acids (EAA) in autumn than in the spring. Also, the FAA contents in eggs and nauplii were higher in autumn than in spring. In contrast, the composition of EAA in eggs was constant between seasons, indicating maternal regulation. The highest cumulative egg production was correlated with a high similarity in the free pool of EAA in the food suspension and the female copepod. Thus, the data support the hypothesis that similarity in the free pool of EAA of diet and female promotes high fecundity and egg hatching success in C. finmarchicus.     

“Good” and “bad” diatoms: development,growth and juvenile mortality of the copepod <Emphasis Type="Italic">Temora longicornis</Emphasis> on diatom diets

We measured development, growth and juvenile mortality of the common copepod Temora longicornis on 11 different monospecific diatom diets in order to estimate (1) how common the negative effects of diatoms are on the development of this copepod and (2) whether the arrested development is connected to deleterious polyunsaturated aldehydes (PUA) or food nutritional quality. Four diatom species (Thalassiosira weissflogii, Thalassiosira rotula CCMP1647, Leptocylindricus danicus CCPM469 and Skeletonema costatum CCMP1281) supported complete development, whereas development failed in or before metamorphosis on seven diatom species/strains (Chaetoceros affinis CCMP158, C. decipiens CCMP173, C. socialis, T. rotula CCMP1018, Thalassiosira pseudonana CCMP1010 and CCMP1335). However, four out of these seven species were not ingested by nauplii, either due to morphology (Chaetoceros spp.) or large size (T. pseudonana CCMP1010). The growth rate did not correlate with the ingestion rate of PUA, neither with ingestion of food mineral (nitrogen) nor with biochemical (polyunsaturated fatty acids, sterols) components. We show that, although some diatoms are of inferior food quality, this is unlikely to be connected to toxicity or due to a direct limitation by a single food nutritional compound.     

Hatching and viability of copepod eggs at two stages of embryological development: anoxic/hypoxic effect

The effects of low oxygen concentrations on the hatching and viability of copepod eggs at two stages of embryological development were investigated. Fully developed eggs from Acartia tonsa (Dana) and Labidocera aestiva (Wheeler) collected between July and September 1991 at Turkey Point, Florida, USA, hatched at lower oxygen concentrations than newly spawned eggs given the same incubation periods. Since many of the newly spawned eggs subsequently hatched when exposed to normoxic conditions, it is likely that the exposure to low oxygen delayed embryonic development. At oxygen concentrations where no initial hatching occurred, the subsequent hatching success of fully developed eggs of A. tonsa was higher than that of newly spawned eggs, indicating that newly spawned eggs were more sensitive to low oxygen concentrations. No such difference was observed for the eggs of L. aestiva.     

Ontogenetic feeding shifts in the meiobenthic harpacticoid copepod Nitocra lacustris

¹⁴C-radiolabelling experiments indicate that adult stages of the salt-marsh harpacticoid copepod Nitocra lacustris (Schmankevitsch) receive a large part of their nutrition through the ingestion and assimilation of certain diatoms. An abundance of empty diatom frustules occurs in the gutpellet contents of field-collected individuals. Naupliar stages do not ingest diatoms in the laboratory, and nauplii from the field do not contain frustules in their gut pellets. Ingestion of diatoms in the laboratory first occurs during the second or third copepodite stage. ³H-radiolabel expeiments and grazing experiments using bacterium-sized beads adhering to the diatoms indicated that both adults and nauplii ingest bacteria adhering to the outer mucus coating of the diatoms (and probably ingest the diatom mucus itself). Adults ingest bacteria (and probably mucus exopolymer) coincidently while ingesting diatoms. The nauplii ingest these components by scraping the outer surface of the diatoms. SEM observations indicate that diatoms are not punctured by the nauplii during feeding. While diatom mucus and associated bacteria play an (as yet unquantified) role in the nutrition of the adults, these components may comprise the bulk of food resources for naupliar stages.     

Egg production and hatching success in North Adriatic Sea populations of the copepod acartia clausi

Egg production, fecal pellet production and hatching success are reported for Acartia clausi females sampled during three cruises in February 1997, 1998 and June 1997 at 20-24 stations along 4 transects in the North Adriatic Sea. Dramatically low hatching rates were recorded during both diatom bloom events in February as opposed to much higher rates during post-bloom conditions in June, even though A. clausi productivity during the bloom was apparently high. These results are discussed in the light of recent findings on the negative impact of diatoms on copepod reproductive potential.     

The effects of temperature and salinity on egg production and hatching success of Baltic Acartia tonsa (Copepoda: Calanoida): a laboratory investigation

The functional response of the aspects of reproductive success of a southwestern Baltic population of Acartia tonsa (Copepoda: Calanoida) was quantified in the laboratory using wide ranges in temperatures and salinities. Specifically, daily egg production (EP, # female⁻¹ day⁻¹) was determined for 4 or 5 days at 18 different temperatures between 5 and 34°C and the time course and success of hatching were evaluated at 10 different temperatures between 5 and 23°C. The effect of salinity (0 to 34 psu) on egg hatching success was also examined. The highest mean rates of EP were observed between 22 and 23°C (46.8–50.9 eggs female⁻¹ day⁻¹). When studied at 18 psu, hatching success of eggs increased with increasing temperature and was highest (92.2%) at 23°C. No hatching was observed for eggs incubated at low temperatures (≤12°C) that were produced by females acclimated to temperatures ≤10°C indicating a possible thermal threshold between 10.0 and 13.0°C below which only the production of diapause (or low quality) eggs exists in this population. When tested at 18°C, the hatching success of eggs incubated at 15 different salinities increased asymptotically with increasing salinity and was maximal (81.4–84.5%) between 17 and 25 psu. The high reproductive success observed over wide ranges in temperatures and salinities in this Baltic population demonstrates one of the mechanisms responsible for the cosmopolitan distribution of this species within productive, estuarine and marine habitats.     

Lethal and sublethal effects of naphthalene and 1,2-dimethylnaphthalene on the marine copepod <Emphasis Type="Italic">Paracartia grani</Emphasis>

Here we evaluate the effects of two quantitatively very important components of the water soluble fractions of fuel oils (naphthalene and 1,2-dimethylnaphthalene, hereafter NAPH and C2-NAPH, respectively) on the survival, feeding and egg production rates, and viability of eggs of the coastal copepod Paracartia (Acartia) grani. Acute toxicity responses resulted in lethal concentrations (LC50) of 2,535 and 161 μg l⁻¹ for NAPH and C2-NAPH, respectively, with no evidence of narcotic effects. Hydrocarbon-specific differences in the toxicity response indicate that sublethal effects (EC50) on feeding by C2-NAPH were likely driven by induced mortality, whereas NAPH has direct negative effects on feeding. Sublethal effects on egg production rates followed a similar detrimental pattern to the one exhibited by feeding rates, suggesting that the lower egg production rates were mediated by the decrease in feeding rates. At the exposure time tested (24 h), the 50% reduction effective concentrations (EC50) determined for sublethal effects were relatively high in comparison with hydrocarbons’ concentrations found under natural circumstances. Long exposure (4 days) of P. grani adults to the tested hydrocarbons at concentrations well below the recorded EC50, however, had no significant effects on feeding, egg production and hatching rates. The viability of the eggs was either not affected or only slightly influenced when healthy eggs were incubated under very high concentrations (up to 6,400 and 700 μg l⁻¹ NAPH and C2-NAPH, respectively). The significance of the effects of oil spills on marine zooplankton communities is discussed in light of the results presented in this study.     

EGG PRODUCTION AND HATCHING SUCCESS IN NORTH ADRIATIC SEA POPULATIONS OF THE COPEPOD Acartia clausi

Egg production, fecal pellet production and hatching success are reported for Acartia clausi females sampled during three cruises in February 1997, 1998 and June 1997 at 20-24 stations along 4 transects in the North Adriatic Sea. Dramatically low hatching rates were recorded during both diatom bloom events in February as opposed to much higher rates during post-bloom conditions in June, even though A. clausi productivity during the bloom was apparently high. These results are discussed in the light of recent findings on the negative impact of diatoms on copepod reproductive potential.     

Laboratory studies of the marine copepod Centropages typicus: egg production and development rates

Egg production and development rates of Centropages typicus (Krøyer) were studied in the laboratory under carying food and temperature conditions. Egg production rates in the laboratory ranged from 0 to 124 eggs female^-1 d^-1 and increased with food concentration up to a critical food concentration (P_c) above which egg production was constant. Egg production rates were influenced by temperature, with more eggs being produced at 15°C than at 10°C. Thalassiosira weisflogii and Prorocentrum micans were determined to be equally capable of supporting egg production at concentrations above P_c at 15°C. Rate of egg production was independent of adult female size when food and temperature were constant. Egg production rates of freshly captured females ranged from 0 to 188 eggs female^-1 d^-1 and were higher in April and May than in June or July. Hatching rates of eggs increased with increased temperature; 95% of the eggs at 15°C hatched within 48 h, while only 8% of the eggs at 10°C hatched within 48 h. Development rates, determined at 10°C in excess concentrations of T. weisflogii, were 23.0 d from egg release to copepodid state I, 27.0 d to stage II, 29.5 d to stage III, 32.2 d to stage IV, 38.5 d to stage V and 49 d to adulthood based on the average time required for 50% of the organisms in an experiment to attain a given stage. Adult males were usually observed 2 to 4 d before adult females, and therefore have a slightly faster rate of development. The effects of temperature, food type and food concentration on egg production and the seasonal appearances of diatoms in the New York Bight may account for the observed seasonal cycles in abundance of C. typicus in these coastal waters.