Effects of anoxia on the viability of subitaneous eggs of planktonic copepods

The effect of anoxia on the viability (as evidenced by hatching) of newly spawned and fully developed subitaneous eggs of three species of copepods, Acartia tonsa (Dana), Centropages hamatus (Lilljeborg), and Labidocera aestiva (Wheeler) was determined for eggs produced in the laboratory by adults collected from inshore waters of the northeastern Gulf of Mexico between January and August 1992. Hatching success decreased to 50% or less when eggs were exposed to oxygen concentrations <0.06 mll^-1 for more than 12 d, except for newly spawned eggs of L. aestiva, which still showed 50% survival after 32 d of exposure to anoxia. For all three species, newly spawned eggs survived exposure to anoxia longer than fully developed eggs. The results indicate that the increasing occurrence of anoxia in estuarine and coastal waters could have a major impact on the population growth of these important food web organisms.     

Wave action: the environmental trigger for hatching in the California grunion Leuresthes tenuis (Teleostei: Atherinopsidae)

 California grunion Leuresthes tenuis (Teleostei: Atherinopsidae) emerge from the ocean to spawn on beaches of southern California. Grunion eggs do not hatch at a set developmental age, but remain in the sand until turbulent surf at high tide washes them out to sea. In previously studied fishes and amphibians that lay eggs terrestrially, low oxygen is the trigger for hatching in water, and high oxygen tensions inhibit hatching. For the grunion, however, eggs placed in air, seawater and oil did not hatch at any at oxygen tension until they were agitated in fluid. Following agitation in seawater, all eggs hatched within several minutes. Grunion eggs in normoxic or hyperoxic water hatched significantly faster than eggs agitated at the lowest oxygen tensions. Mechanical agitation, not hypoxia, is the environmental trigger for hatching in California grunion eggs. Received: 15 May 1999 / Accepted: 5 April 2000     

Effects of various environmental factors on egg development of several species of Acartia in Southern California

The hatching of the eggs of 3 Acartia species (A. tonsa, A. clausi, and Acartia sp. I) spawned in the laboratory and the eggs of Acartia (species undetermined) isolated from natural bottom sediments was examined under various environmental conditions (temperature, salinity and temperature combinations, oxygen concentration, illumination, and the presence of bottom mud). The range of temperature and salinity in which eggs hatched differed from one species to another. Egg hatching was inhibited under certain environmental conditions for all species, although egg viability under such conditions varied from one species to another. In relation to these findings, habitat separation of planktonic Acartia populations and the role of dormant eggs during their seasonal absence in the plankton of Southern California coastal waters are discussed.     

Effects of low oxygen concentrations on the hatching and viability of eggs of marine calanoid copepods

Hatching of the eggs of four species of copepods, Acartia tonsa, Labidocera aestiva, Tortanus discaudatus, and Centropages hamatus was suppressed at oxygen concentrations less than 0.02 ml O₂/l. When such eggs were subsequently exposed to normal oxygen concentrations, hatching varied among the species, indicating variability in the capacity of eggs to survive exposure to low oxygen. Incubation of eggs at low oxygen concentrations resulted in an increase in the duration of embryonic development. Experiments were conducted in closed systems and oxygen concentration was determined by thermal conductivity using gas chromatography.     

Potential effects of a resuspension event on the vertical distribution of copepod eggs in the sea bed: a laboratory simulation

Adult female Acartia tonsa and Labidocera aestiva were obtained from inshore marine waters near Turkey Point, Florida (29°54.5N;84°31W) during April and May 1991. For each species, eggs spawned overnight in the laboratory were collected, mixed with sediment, and added to a plastic tube (26 to 29 cm longx5 cm diam) to yield a 2 cm-thick layer. This tube was connected to a 245 cm-long tube, seawater was added and the contents were thoroughly mixed by turning the assembly end-over-end several times. The vertical distribution of eggs in the sediments following resuspension varied according to the grain size of the sediments. For sediments of 125 m particle-diameter, 68 to 72%, and 70 to 73% of the eggs of A. tonsa and L. aestiva, respectively, occurred in the upper 4 mm.For sediments 63 to <125 m particle-diameter, 34 to 36% of the eggs of A. tonsa and L. aestiva, respectively, occurred in the upper 4 mm. For sediments of <63 m particle-diameter, egg distribution was uniform for both species throughout the 2 cm layer. The results indicate that the vertical distribution of eggs in the sea bed following a resuspension event should be highly dependent upon the grain-size composition of the sediments.     

Hydrostatic pressure-induced apoptosis on nauplii of Calanus sinicus

The effect of hydrostatic pressure on embryonic development of the calanoid copepod Calanus sinicus was studied. Differences of pressure effect among blastomere stages, 1-cell, 2-cell, 4-cell, 8-cell, 16-cell, blastula and limb-bud stage, were examined under two pressurizing conditions, abruptly (10 atm/min) and gradually (0.1 atm/min) increasing. Egg hatching success, deformity frequency and apoptotic cell degradation of hatched nauplii were examined. Egg hatching success rate was not significantly different between blastomere stages, and also between pressurizing conditions. Deformity frequencies of hatched nauplii were low in the early 1-cell and 2-cell stages, and high in the later blastula and limb-bud stages, in both abrupt and gradual pressurizing conditions. On the other hand, in regard to difference in pressurizing conditions, deformity frequency in gradual pressurizing was significantly higher than that in abrupt pressurizing rate. Gradual pressure increase seems to be more harmful to C. sinicus eggs than abrupt pressure change. Apoptosis induced in nauplii by hydrostatic pressure was detected for the first time in marine zooplankton. The embryos of C. sinicus are sensitive to pressure variations, that is, these embryos are supposed to sink to deeper waters, incurring greater risk of having deformities. In the field, C. sinicus ascend to the surface and spawn at night. By looking from this upward behavior, eggs are spawned at lower pressure and warmer temperature. Probably, the harmless low pressure and warm temperature lead eggs to hatch early and to recruit successfully.     

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.     

Distribution and abundance of eggs of Labidocera aestiva (Copepoda: Calanoida) in the bottom sediments of Buzzards Bay,Massachusetts, USA

Grain-size composition of the sea-bed and density of eggs of Labidocera aestiva in bottom sediments in Buzzards Bay were determined at approximately monthly intervals from March 1983 through April 1984. The results of this study, together with those of Marcus (1984), show that during the fall and winter periods of 1982–1984 the proportion of eggs of L. aestiva occurring in the surficial sea-bottom sediments declined and the proportion of eggs in the deeper sediment layers increased. High positive correlations (r ²=0.72 and 0.92) were observed during the early fall 1983 between egg abundance and the proportion of the total sediments represented by the mud fraction. During late fall, winter, and early spring 1983–1984, the coefficients of determination were much lower. Physical criteria (e.g. sedimentation and transport characteristics) can be used to accurately predict the distribution and abundance of recently spawned eggs on the sea-bottom.     

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.     

Diel,lunar and seasonal rhythms in the reproduction of two tropical damselfishes: Pomacentrus flavicauda and P. wardi

It has been hypothesized that site-attached coral reef fishes breed during periods when their offspring will be rapidly flushed from the reef environment, but within the season when these propagules are least likely to be forever lost from the local populations. Daily, monthly and annual rhythms were observed in the reproduction between November 1978 and February 1980 of two tropical damselfishes, Pomacentrus flavicauda and P. wardi, from the southern Great Barrier Reef, Australia. These data support the first part of the hypothesis, but not the explanation for seasonality. When breeding, females of both species spawned demersal eggs into the care of males during a brief period at first light. After several days, larvae hatched from these eggs during another brief period after sunset. An experiment confirmed that the change from light to dark triggered the hatchings and that the emerging larvae were photopositive. Eggs of P. flavicauda usually hatched on the evening of their fourth day in the nest; eggs of P. wardi hatched on their fifth day. A small proportion of eggs required longer incubation. Hatching success was 90%. When breeding, both species spawned in bi-weekly cycles synchronised with the phases of the moon. The largest clutches, as well as the most clutches, were laid at the peaks of these cycles. In the case of P. flavicauda, maximum hatchings occurred on days during which spring high tides fell near sunset (the cue for hatching to begin). It is likely that these hatchlings were rapidly flushed off the reef by the ebbing tides. The hatching cycles of P. wardi consistently peaked 4 d earlier in the lunar month. Consequently, hatchlings of this species remained in shallow water at least one tidal cycle longer than those of P. flavicauda. This difference may be a consequence of the different habitat specialisations of the two species. Both species bred seasonally during the warmest, calmest months of the year. This is not the period of least current flow on the Great Barrier Reef. The observed seasonality may reflect the best times for larval survival in the open sea rather than an adaptation that limits the loss of offspring from the parent populations.     

Zooplankton responses to hypoxia: behavioral patterns and survival of three species of calanoid copepods

Seasonally recurrent and persistent hypoxic events in semi-enclosed coastal waters are characterized by bottom-water dissolved oxygen (d.o.) concentrations of < 2.0 ml l⁻¹. Shifts in the distribution patterns of zooplankters in association with these events have been documented, but the mechanisms responsible for these shifts have not been investigated. This study assessed interspecific differences in responses to hypoxia by several species of calanoid copepods common off Turkey Point, Florida, USA: Labidocera aestiva (Wheeler) (a summer/fall species), Acartia tonsa (Dana) (a ubiquitous year-round species), and Centropages hamatus (Lilljeborg) (a winter/spring species). Under conditions of moderate to severe hypoxia 24-h survival experiments were conducted for adults and nauplii of these species from August 1994 to October 1995. Experiments on adults used a flow-through system to maintain constant d.o. concentrations. Adults of A. tonsa showed no decline in survival with d.o. as low as 1.0 ml l⁻¹, sharp declines in survival at d.o. = 0.9 to 0.6 ml l⁻¹, and 100% mortality with d.o. = 0.5 ml l⁻¹. Adults of L. aestiva and C. hamatus were more sensitive to oxygen depletion: both species experienced significant decreases in survival for d.o. = 1.0 ml l⁻¹. Nauplii of L. aestiva and A. tonsa showed no significant mortality with d.o. = 1.1 to 1.5 ml␣l⁻¹ and d.o. = 0.24 to 0.5 ml l⁻¹, respectively. In addition, experiments investigating behavioral avoidance of moderate to severe hypoxia were carried out for adults of all three species. None of the three species effectively avoided either severely hypoxic (d.o. < 0.5 ml l⁻¹) or moderately hypoxic (d.o. ≈ 1.0 ml l⁻¹) bottom layers in stratified columns. These results suggest that in␣nearshore areas where development of zones of d.o. < 1.0 ml l⁻¹ may be sudden, widespread, or unpredictable, patterns of reduced copepod abundance in bottom waters may be due primarily to mortality rather than avoidance. Received: 31 August 1996 / Accepted: 24 September 1996     

Hydrostatic pressure-induced apoptosis on nauplii of Calanus sinicus

The effect of hydrostatic pressure on embryonic development of the calanoid copepod Calanus sinicus was studied. Differences of pressure effect among blastomere stages, 1-cell, 2-cell, 4-cell, 8-cell, 16-cell, blastula and limb-bud stage, were examined under two pressurizing conditions, abruptly (10 atm/min) and gradually (0.1 atm/min) increasing. Egg hatching success, deformity frequency and apoptotic cell degradation of hatched nauplii were examined. Egg hatching success rate was not significantly different between blastomere stages, and also between pressurizing conditions. Deformity frequencies of hatched nauplii were low in the early 1-cell and 2-cell stages, and high in the later blastula and limb-bud stages, in both abrupt and gradual pressurizing conditions. On the other hand, in regard to difference in pressurizing conditions, deformity frequency in gradual pressurizing was significantly higher than that in abrupt pressurizing rate. Gradual pressure increase seems to be more harmful to C. sinicus eggs than abrupt pressure change. Apoptosis induced in nauplii by hydrostatic pressure was detected for the first time in marine zooplankton. The embryos of C. sinicus are sensitive to pressure variations, that is, these embryos are supposed to sink to deeper waters, incurring greater risk of having deformities. In the field, C. sinicus ascend to the surface and spawn at night. By looking from this upward behavior, eggs are spawned at lower pressure and warmer temperature. Probably, the harmless low pressure and warm temperature lead eggs to hatch early and to recruit successfully.     

Deposit-feeding amphipods (Monoporeia affinis) reduce the recruitment of copepod nauplii from benthic resting eggs in the northern Baltic Sea

We experimentally investigated the effect of different densities of the burrowing, deposit-feeding amphipod Monoporeia affinis on the recruitment of zooplankton from benthic resting eggs. Intact sediment cores with in situ density and species composition of zooplankton resting eggs and benthic fauna were collected in the northern Bothnian Sea, part of the Baltic Sea. We removed as many M. affinis as possible from the cores and then added different numbers of M. affinis to the cores to generate a range of densities. The cores were exposed to different densities of M. affinis for either 3 or 40 days, after which the hatched zooplankton was registered. One subset of the cores were initially incubated under low temperature (2–3 °C, to prevent hatching) for 37 days (the resting phase), to allow for effects of M. affinis on unhatched resting eggs. These cores were then incubated under higher temperature (13 °C) for 3 days (the hatching phase), to induce hatching and allow for effects on hatching or hatched specimens. In a second subset of cores with the same time and temperature schedule, the M. affinis density was experimentally reduced at the start of the hatching phase, to evaluate the effect of M. affinis during the hatching phase. To a third subset of cores, we immediately initiated the hatching phase, without an experimental resting phase, to evaluate the effects induced during the resting phase. The most common zooplankton species that hatched was Eurytemora affinis (Copepoda), followed by Bosmina longispina maritima (Cladocera). In all cores that were subjected to a resting phase, the numbers of hatched E. affinis were log-linearly negatively related to density of M. affinis. An increase of M. affinis density from 1,000 to 5,000 individuals m⁻², normal field densities, reduced the hatching by 60–70%. The negative impact was mainly exerted during the hatching phase, suggesting predation on, burial of or physical injury of hatching nauplii or eggs in a late development stage as likely mechanisms. Also, the number of B. longispina maritima that hatched was reduced by M. affinis during the hatching phase, but no clear relation to density of M. affinis could be identified. The results show that M. affinis can reduce recruitment to zooplankton from benthic resting eggs. Such impact by the benthos on resting stages of zooplankton is therefore a potentially significant link between the benthic and pelagic systems. Received: 10 August 2000 / Accepted: 13 November 2000     

Mating,egg-laying and hatching of Octopus tetricus (Mollusca: Cephalopoda) in the laboratory

Observations on mating, egg-laying and hatching of aquarium-held individuals of Octopus tetricus Gould, 1852 are described. Females were able to store viable spermatozoa for periods up to 114 days, and laid many small eggs in strings 10 to 12 cm long. The duration of embryonic development varied inversely with temperature. The newly hatched planktonic larvae were 2.5 mm long by 1.1 mm wide, but attempts to rear them were unsuccessful, the longest survival being 21 days. The females died following egg hatching.     

Carbofuran induced changes in breeding of a freshwater fish,Labeo rohita (Hamilton)

The effect of carbofuran, a carbamate pesticide, on the breeding performance of the freshwater fish Labeo rohita (Hamilton) has been investigated in the present study. Breeding of L. rohita was conducted after treatment with three sublethal concentrations, i.e. 0.06, 0.15 and 0.30 mg L^?1 of carbofuran for 96 h. The present investigation showed that the number of total eggs and total amount of eggs (litre per kg body weight) decreased significantly (p < 0.01) at all concentrations of carbofuran in comparison to controls, while the reduction in fertilization percentage at all concentrations of carbofuran was not significantly different from controls. The reduction in hatching percentage, the expected number of hatchlings, and the expected number of hatched larvae were significantly (p < 0.01) different between treated and control groups at all carbofuran concentrations. No significant differences for the 96 h survivability of hatched larvae were reported at all concentrations of carbofuran.     

Growth, carbon, nitrogen and caloric content of Solea senegalensis (Pisces: Soleidae) from egg fertilization to metamorphosis

Eggs and larvae of the Senegal sole, Solea senegalensis Kaup, were reared from fertilization until the end of metamorphosis, which occurs by Day 17 after hatching at 19.5 °C. Changes in energy content and biomass quality were studied in terms of dry weight and of carbon, nitrogen and energy content. S. senegalensis spawned eggs of about 1 mm diameter which hatched 38 h after fertilization. Average dry weight of individual eggs was 46 μg, the chorion accounting for about 18% of total dry weight. Gross energy of recently fertilized sole eggs was approximately 1 J egg⁻¹. From fertilization to hatching, eggs lost 8% of their total energy (chorion not included). After hatching, larvae lost 14% of their initial energy until the start of feeding which occurred about 48 h afterwards. The principal components catabolized during embryogenesis were carbon-rich compounds that decreased by 26%, while nitrogen-rich compounds decreased by only 10% and were practically unaltered from hatching to the start of feeding. Feeding larvae displayed constant growth during the period studied (specific growth rate on a dry weight basis was 0.26 d⁻¹). The relative proportion of carbon and nitrogen content revealed an accumulation of high energy compounds in the days before metamorphosis. By Day 14, the energy content reached values similar to those of recently hatched embryos, but decreased again during metamorphosis. Received: 10 June 1998 / Accepted: 28 January 1999     

Poecilogony in an estuarine opisthobranch: planktotrophy, lecithotrophy, and mixed clutches in a population of the ascoglossan Alderia modesta

A San Diego population of the opisthobranch mollusc Alderia modesta (Lovén, 1844) exhibits poecilogony, the presence of two development modes within a single species. In spring, half of the adults spawned masses containing ∼300 eggs with a mean diameter of 68 μm. After 3 d, these egg masses hatched planktotrophic veligers with a maximum shell dimension of 116 μm. The remaining adults spawned masses containing ∼30 eggs with a mean diameter of 105 μm. These egg masses hatched after 5 to 6 d, releasing lecithotrophic larvae with a maximum shell dimension of 186 μm. About 1% of field-collected adults produced mixed clutches containing a continuum of larval sizes, spanning the size extremes of planktotrophy and lecithotrophy and hatching larvae with a mean maximum shell dimension of 152 μm. Adults producing planktotrophic and lecithotrophic larvae were interfertile, and no hybrid breakdown was observed through the F3 generation. When starved, adults which previously produced only lecithotrophic larvae switched to producing planktotrophic larvae or mixed clutches with both planktotrophic and lecithotrophic larvae. Sequence-polymorphisms from a portion of the mitochondrial cytochrome c oxidase I gene support the conclusion that the two reproductive morphs represent a single species. Most of the lecithotrophic larvae and a small percentage of the larvae from mixed clutches were metamorphically competent within 3 d of hatching. A. modesta is the only molluscan species as yet known to have both planktotrophic and pelagic lecithotrophic development within a single natural population. Received: 14 August 1997 / Accepted: 11 April 1998     

Effects of anoxia and hypoxia on the dormancy and survival of subitaneous eggs of<Emphasis Type="Italic"> Acartia bifilosa</Emphasis> (Copepoda: Calanoida)

Subitaneous eggs of the copepod Acartia bifilosa were incubated at low oxygen concentrations and in anoxic conditions to study the induction of dormancy and the longevity of the resting eggs. Eggs were incubated in hypoxic conditions for 8 or 10 days, and also in anoxic conditions for 8–441 days. After the anoxic and hypoxic incubations, the unhatched eggs were subsequently incubated in normal oxygen conditions to reveal the viability of the eggs. Low oxygen concentrations (< 0.17 ml l^–1) induced dormancy in the eggs: they did not hatch at these concentrations in the hypox experiments but did hatch afterwards. Eggs survived well in anoxic conditions: after 10 months at 4°C, approximately 40% of the eggs still hatched, while some eggs survived in anoxic conditions for a year at 13°C, which is longer than hitherto reported for subitaneous eggs of any copepod species. Also, sediment that contained eggs of copepods was stored in the laboratory to reveal the long-term survival of the eggs. When incubated after 1.8 years storage, more than 40% of A. bifilosa eggs hatched. The study shows that subitaneous eggs may also be used as a survival strategy on seasonal time scales. This may partly explain why A. bifilosa does not produce diapause eggs in the present study area in the northern Baltic Sea.Communicated by M. Kühl, Helsingør     

Calanoid copepod eggs in sea-bottom muds. V. Seasonal changes in hatching of subitaneous and diapause eggs of Tortanus forcipatus

A warm-water neritic calanoid copepod, Tortanus forcipatus, has been found to lay two physiologically different types of eggs (subitaneous and diapause eggs), which are separately spawned in response to different seasonal environmental conditions. Subitaneous eggs are produced when the planktonic populations are at their numerical maxima, and are only shed until late summer. After peak spawning, true resting eggs are spawned which undergo diapause for 1 to 3 months. The eggs recover from their diapause around mid-winter, but hatching does not take place until the water temperature at the sea bottom exceeds 15°C in early summer.     

Reproduction of the Antarctic fish Nototheniops nudifrons

The reproductive biology of the Antarctic fish Nototheniops nudifrons (Lönnberg, 1905) was analyzed by examination of the gonads of fish collected in March and April 1985 in trawls near Low Island, Antarctic Peninsula, and compared to direct observations of reproduction and early larval development in the laboratory from March to October of the same year. Males and females reached sexual maturity at an age of 4 to 5 yr. Mature males differed in coloration from females and immature males. Ovaries of sexually mature females contained two distinct size modes of vitellogenic oocytes, representing two separate clutches of developing eggs. Females spawned 100 to 3 500 demersal eggs, which were laid in a nest in crevices or under rocks, and guarded by the male for about 4 mo. Females did not assist in nest defense or egg care. Most spawning in the field and in the laboratory occurred in late fall and early winter (May to June). A second clutch may be spawned in spring (November and December). Eggs hatched after 124 d, and larvae were raised for 38 d. Otoliths of larvae contain internal microincrements, which are deposited in a daily fashion, and are visible by light microscopy. The otoliths of 32 adult fish were examined by scanning electron microscope, and counts of microincrements in these otoliths allowed the backcalculation of hatching dates. Estimated hatching dates were between September and May.