Selective accumulation of mycosporine-like amino acids in ovaries of the green sea urchin Strongylocentrotus droebachiensis is not affected by ultraviolet radiation

Field sampling and laboratory experiments examined whether ultraviolet radiation (UVR) affects the reproduction or the accumulation of mycosporine-like amino acids (MAAs) and ascorbic acid in ovaries of the green sea urchin Strongylocentrotus droebachiensis (Müller). Ovaries of sea urchins sampled across a depth gradient (0.5–10 m) in March 1998 did not differ in their gonadal index, or in concentrations of MAAs, or ascorbic acid. Concentrations of MAAs and ascorbic acid in ovaries were higher in sea urchins collected from a kelp bed compared with those collected from a community of crustose coralline algae. The concentrations of MAAs in ovaries varied seasonally, peaking in March, when sea urchins had high gonadal indices just before spawning. Ovaries of sea urchins maintained on controlled diets from October 1997 to April 1998 accumulated significantly higher concentrations of MAAs when fed a diet rich in MAAs than did ovaries of sea urchins fed an alga lacking MAAs, but the gonadal indices did not differ between diets. Sea urchins accumulated principally one MAA, shinorine, but not others that were available in high concentrations in their diet. Neither the gonadal index nor the ovarian concentrations of MAAs were affected by daily exposure of adult urchins to UVR for 6 months. Concentrations of ascorbic acid in ovaries did not differ among diets or UV-treatments. The percentages of nutritive phagocytes and gametic cells were not affected by diet or UVR, and did not co-vary with concentrations of MAAs or ascorbic acid in ovaries. These data support previous demonstrations that female sea urchins accumulate MAAs from their diet of macroalgae, but further show that the accumulation is selective for specific MAAs, particularly shinorine, and that adult S. droebachiensis do not accumulate MAAs in their ovaries or eggs in response to UV-exposure. These are also the first experimental studies to address whether MAAs are affected by or regulate gametogenesis, and indicate that they do not. Received: 5 May 2000 / Accepted: 29 September 2000     

Size-dependent reproduction in the Caribbean gorgonian Pseudoplexaura porosa

Reproductive success among clonal taxa is often portrayed as a simple function of clone size, but reproduction in Pseudoplexaura porosa (Houtuyn), a common Caribbean gorgonian, reflects a more complex relationship between size and gamete production. Tagged colonies were sampled at two reefs in the San Blas Islands, Panama during the 1995 spawning season to determine the colony size at first reproduction, size-dependent polyp fecundities, and whole colony gonad production, and to follow the gametogenic cycle. Additional data were obtained during non-spawning months in 1984 and 1985. Of 120 colonies ranging in height from 20 to 250 cm, only colonies taller than 50 cm were reproductive. An average of 4.33 oocytes polyp⁻¹, 560 to 800 μm in diameter, were released during monthly synchronous spawning events from June to September. Oocytes released each month developed from a group of intermediate size oocytes present at the end of the preceding month. Almost all oocytes >560 μm were released during each spawning event. Spermatogenesis was a shorter process than oogenesis. During each spawning month, mature spermaries developed from a group of <125 μm spermaries. Virtually all spermaries >190 μm were released each month, and most of the spermary volume in mature male polyps was generated anew each month of spawning. Among reproductive colonies, polyp fecundity increased with colony height from 50 to 200 cm. Per capita fecundity of >200 cm tall female colonies was lower than for 150 to 200 cm colonies, but whole colony fecundity of large colonies was greater due to the exponential increase in the number of polyps that occurs with increasing colony height. In male colonies, both polyp and colony reproductive output increased with colony height. The large amounts of sperm produced by large male colonies probably contributes to the high rates of in situ fertilization observed in P. porosa. Differences in fecundity as a function of colony size and sex suggest differences in some combination of the cost of reproduction and/or allocation of resources to reproduction. Received: 9 September 1998 / Accepted: 7 June 1999     

Density-related reproductive trade-offs in the green sea urchin, Strongylocentrotus droebachiensis

Green sea urchin, Strongylocentrotus droebachiensis (O.F. Müller), populations are being depleted rapidly in the Gulf of Maine and there is justified concern that the potential of this free-spawner to produce larvae may be severely inhibited. We evaluated the opposing effects of different population densities on gonad development and fertilization success, using population surveys and fertilization experiments. We determined gonad indices (gonad mass/body mass) over a range of population densities (0.1 to 250 ind. m⁻²) at seven sites in coastal Maine, USA, sampled at two depths (5 and 15 m). At shallow sites, we found that gonad indices declined by 50% over the 1500-fold range in adult population density. At 15 m deep locations, gonad mass was consistently low and did not vary significantly with density. Patterns of macroalgal abundance suggest urchins at high density and in deeper water were food limited. Because macroalgal cover co-varies inversely with sea urchin density, we designed field experiments to determine the interaction between sea urchin density and kelp canopy on fertilization success. On square arrays we manipulated the spacing of simulated urchins, but held their numbers constant (five sperm-filled syringes interspersed with four Nitex mesh egg containers permeable to sperm). These experiments, simulating the observed range of natural density, suggested that (1) fertilization rates decreased many times faster than individual gamete production increased over the same range in density, and (2) kelp increased fertilization success at high density when eggs were within 25 cm of a sperm source, but not when spaced 1 m apart. Additional laboratory fertilization experiments at ambient temperatures (3 to 5 °C) indicated that diluted sperm were viable for <1 h, but egg viability was virtually unchanged for >8 h. In short, to the individual the reproductive benefits of aggregating appear to outweigh the costs; and while sperm may be limiting at low population density, eggs may remain viable long enough to be fertilized by sperm from more distant males. Received: 10 February 1998 / Accepted: 22 January 1999     

Water-soluble extracts of the diatom Thalassiosira rotula induce aberrations in embryonic tubulin organisation of the sea urchin Paracentrotus lividus

Eggs and embryos of the sea urchin Paracentrotus lividus were used as a model to study the effect at the cellular level of potential anti-mitotic compounds extracted from the diatom Thalassiosira rotula. Eggs and embryos incubated in a water-soluble diatom extract, corresponding to 5 × 10⁶ and 10⁷ cells ml⁻¹, were totally blocked (i.e. cell division was blocked) at the one-cell stage. At lower concentrations (2.5 and 1.25 × 10⁶ cells ml⁻¹), the first mitotic division was inhibited in 32 ± 26% and 25 ± 3.5% of the zygotes, respectively, demonstrating the dose-dependent effect of diatom extracts on sea urchin development. Immunofluorescence dyes, specific for DNA and α-tubulin subunits, were used to stain nuclei and microtubules in sea urchin embryos during various phases of development. Images with the confocal laser scanning microscope showed that tubulin was not organised in filaments at the sperm aster and cortex levels, and that the pronuclei were not fused in embryos incubated soon after fertilisation with water-soluble diatom extracts corresponding to 10⁷ cells ml⁻¹. At lower diatom-extract concentrations (4 × 10⁶ cells ml ⁻¹), fusion of the pronuclei occurred but the mitotic spindle was not formed. Microtubules were clearly de-polymerised and the chromatin appeared globular and compacted at the centre of the cell. A similar structure was observed for sea urchin embryos incubated with 0.1 mM colchicine, a potent anti-mitotic compound. When sea urchin embryos were incubated in water-soluble diatom extracts at different times prior to the first mitotic division, microtubules appeared de-polymerised at each step, from pronuclear fusion to telophase, and cell division was blocked. At the histological level, embryos incubated with 4 × 10⁶ cells ml⁻¹ diatom extract showed nuclear fragmentation without cytokinesis. The possible use of sea urchin embryos as a bioassay to test for other unknown compounds with cytotoxic activity in phytoplankton species is discussed. Received: 7 May 1998 / Accepted: 9 December 1998     

Sperm precedence and sperm storage in multiply mated red flour beetles

In insects, the last male to mate with a female often gains access to a disproportionate number of subsequent fertilizations. This study examined last-male sperm precedence patterns in doubly and triply mated Tribolium castaneum females. Sperm storage processes were investigated by measuring the quantity of sperm stored within the female spermatheca following single, double, and triple matings. Both doubly mated and triply mated females exhibited high last-male sperm precedence for progeny produced during the first 48 h following the last mating, with females in both groups exhibiting parallel declines in sperm precedence 1 and 2 weeks later. The number of sperm stored by females increased by 33% between singly mated and doubly mated females, indicating that the spermatheca is filled to only two-thirds capacity following insemination by the first male. Based on the proportion of stored sperm from first and second matings, we tested predictions about sperm precedence values based on models of random sperm mixing. High initial last-male sperm precedence strongly supports stratification of last-male sperm. By 1–2 weeks after double matings, sperm precedence declined to levels indistinguishable from values expected under random mixing. These results provide insight into mechanisms of sperm storage and utilization in this species. Received: 20 August 1997 / Accepted after revision: 24 May 1998     

Sperm economy and limitation in spiny lobsters

Sperm limitation, when female fertilisation success is constrained by the supply of sperm, is generally perceived to be an uncommon feature of reproduction in species which directly transfer gametes during copulation. Male size, previous copulations, and the balance of expected reproductive return and future mating opportunity may, however, limit the amount of sperm males transfer to females. We used laboratory experiments where mate size could be manipulated and its consequences on spermatophore size and clutch size determined, to show that in two genera of spiny lobsters (Crustacea: Palinuridae) male reproductive output limits the size of clutches brooded by females. In Panulirus argus from the Florida Keys, we show that while male size affects spermatophore area, males also vary the amount of ejaculate positively with female size. Furthermore, the area of the spermatophore has a greater influence than female size on subsequent clutch weight. In Jasus edwardsii from New Zealand, female size, male size and mate order all affect clutch weight. In both species, clutches fertilised by small males in the laboratory are significantly smaller than clutches fertilised by large males. These results suggest that to ensure they receive sufficient sperm, females should either mate several times prior to oviposition, mate as early as possible in the reproductive season, or choose large, preferably unmated males as partners and thus compete with other females for preferred males. Sperm-limited female fecundity has the potential to limit the egg production of fished populations where large males are typically rare. Received: 18 May 1998 / Received in revised form: 20 November 1998 / Accepted: 30 November 1998     

Manipulation of food and photoperiod promotes out-of-season gametogenesis in the green sea urchin, Strongylocentrotus droebachiensis: implications for aquaculture

Green sea urchins, Strongylocentrotus droebachiensis OF Müller, collected off the coast of New Hampshire, USA, in late February 1995, were brought into the laboratory and fed an artificial diet ad libitum, and subjected to a photoperiod advanced by 4 months. During this study, temperatures and salinities for experimental urchins mirrored those recorded at the collection site. We examined the effects of changes in feeding regime and photoperiod on gametogenesis and compared the experimental urchins with those from the source population. During the 7-month period, experimental urchins showed no detectable changes in mean test height or diameter. Experimental urchins had a significantly higher gonad index (GI) in March, April and May (18 ± 6%) compared with field urchins in March (11 ± 3%). Subsequently, experimental urchins had a mean monthly GI of 25 to 30%, while the mean GI for field urchins was 11 to 13%. Gonial cell mitosis and gametogenesis occurred earlier in experimental male and female urchins compared with field urchins. Stereological and histological observations and stage–frequency data showed that the ovaries of experimental urchins were large because of the accelerated development of nutritive phagocytes, the volume fraction (V _v) of which was 89 to 90% of the gonad, while new vitellogenic primary oocytes occupied <1% V _v. In males, stereological and histological observations and stage–frequency data suggested a mobilization of materials from the nutritive phagocytes beginning between June and August, i.e. earlier than in females, and, by September, new gametes occupied a V _v of 49 ± 3% of the testes. Oocyte size–frequency distributions demonstrated that most primary oocytes were <80 μm in diameter between March and September, suggesting that cold temperatures may be needed for completion of vitellogenesis. We describe changes in the two principal cell types in the germinal epithelium of urchin gonads and indicate how knowledge of their population dynamics may be useful in aquaculture applications. Received: 21 February 1997 / Accepted: 3 June 1998     

Mate replacement in experimentally widowed collared flycatchers (Ficedula albicollis): determinants and outcomes

There is increasing evidence that sexual selection may be intense even in socially monogamous birds, resulting from both mate choice and sperm competition. We studied these two modes of sexual selection experimentally by removing paired male collared flycatchers, Ficedula albicollis, from their mates for 2 days and investigating the factors that influenced the likelihood of a replacement male appearing and how the removals influenced paternity. Replacement males (usually neighbouring males) appeared at 81% (n = 37) of nests where males were removed. The likelihood of this appearance was unaffected by the probable reproductive value of the female's clutch to the replacing male. A replacement was, however, less likely when the original male had a large forehead patch, a trait previously shown to be subject to sexual selection in this population. Experimental removal of males increased the level of sperm competition: 74% of experimental broods were multiply sired, compared to 29% of unmanipulated broods in a previous study. Only two factors predicted how paternity was shared between males: removed males fathered more young if removed closer to laying, and if they had larger forehead patches. The former result is consistent with last-male sperm precedence determining paternity, whereas the latter adds to other evidence that forehead patch size is the target of female preference in this species. Our results suggest that females exert some control over male replacement, and also that they may influence the fertilisation success of males by behavioural means. Received: 15 July 1998 / Received in revised form: 16 March 1999 / Accepted: 28 March 1999     

Can synchronous spawning be predicted from environmental parameters? A case study of the lugworm Arenicola marina

The annual epidemic spawning period of a Scottish population of Arenicola marina (L.) has been recorded over a period of 13 yr. This population spawns between mid-October and mid-November in a discrete spawning event over a period of 4 to 5 d. Endocrine manipulation experiments showed that spawning is induced in females only if sufficient titres of PMH (prostomial maturation hormone) are present in the prostomia. These levels are attained during the 2 to 3 wk prior to the natural spawning date. The East Sands, St. Andrews population always spawns during periods of spring tides regardless of tidal amplitude or whether they are full- or new-moon tides. Meteorological data, including sea-temperature data were collected for each year, and correlation of the environmental data with spawning time was attempted. Correlation of spawning times with weather patterns showed that mean daily air pressures were significantly higher during the spawning period than from September to November as a whole. Evidence also suggests that a reduction in sea temperature is required prior to spawning. A significant moderate negative correlation was found between May to July air temperatures and spawning date, suggesting that higher May to July temperatures may induce early spawning. Daily rainfall and wind speed were also lower during the spawning period, but not significantly so. These results indicate that air pressure (or changes therein) may act as a final spawning cue, and the advantages of this are discussed in relation to fertilization success. A model of the interplay between environmental parameters and the endocrine mechanisms controlling the induction of spawning is proposed. Higher than average summer temperatures may advance gametogenesis to bring the population into a state of maturity (full-size oocytes, well-developed sperm morulae), and may also advance spawning time. Once the population has completed gametogenesis, a drop in sea temperature is then required to trigger an increase in endocrine titres within the prostomium, without which spawning cannot be induced by prostomial injection. The population spawns on spring tides; however a lack of clement weather coinciding with the spring tide will result in population-wide spawning being aborted, as in 1996. Clement weather (high pressure, low rainfall and wind speed) in conjunction with spring tides permits spawning to proceed to completion. Received: 21 June 1999 / Accepted: 25 January 2000     

A novel symbiosis between a cyanobacterium, Synechococcus sp., an aplastidic protist, Solenicola setigera, and a diatom, Leptocylindrus mediterraneus, in the open ocean

The abundance, biomass and distribution of Solenicola setigera, a colonial heterotrophic protist found only with the centric chain-forming diatom Leptocylindusmediterraneus, are reported for four major ocean basins. The distribution is cosmopolitan, and abundances and biomass are usually low (<500 colonies l⁻¹); however, in the summer of 1993, we observed a major biomass component (range = 5 to 31 μg C l⁻¹) in the surface waters of the North Atlantic attributable to S. setigera. These colonies of S. setigera were exceptionally large, and unusual in possessing high abundances of Synechococcus sp., a normally solitary cyanobacterium, embedded in the matrix covering the cells. We hypothesize that this relationship was mutually beneficial for both Solenicola setigera and Synechococcus sp. Received: 5 January 1998 / Accepted: 22 May 1998     

Temporal variation in protein, carbohydrate, and lipid concentrations in Paramuricea clavata (Anthozoa, Octocorallia): evidence for summer–autumn feeding constraints

Temporal variations in protein, carbohydrate, and lipid levels were studied in a passive suspension feeder, the gorgonian Paramuricea clavata. The samples were collected every month for mature and immature colonies over a three-year period (1997–2000). The relationship between biochemical composition and reproductive output was examined on the basis of the 1998 and 1999 data. In female and male P. clavata colonies, the tissue displayed differences in lipid concentrations only in winter–spring, due to the high lipid levels attained by female colonies in this period. Immature colonies showed significant differences in lipid concentration only with respect to mature females in spring. There were clear seasonal trends in the lipid and carbohydrate levels in P. clavata, with maximum values in winter–spring [male lipid 212±75 SD μg mg⁻¹ of organic matter (OM) and female lipid 274±103 SD μg mg⁻¹ of OM; male and female carbohydrate 68±14 SD μg mg⁻¹ of OM], coinciding with maximum food concentration/quality, and minimum values in summer–autumn (male and female lipid 155±57 SD μg mg⁻¹ of OM; male and female carbohydrate 56±14 SD μg mg⁻¹ of OM), coinciding with low food concentration/quality. The relationship between reproductive output and tissue concentrations of lipids, carbohydrates, and proteins was not straightforward, although there was an evident overlap of the lipid accumulation and the gonadal development. The results of this study show that protein, carbohydrate, and lipid levels may provide a record of episodes in the ecological cycle bearing on the trophic aspects of the target species. The results indicate that information on seasonal biochemical levels may explain benthopelagic coupling processes, provided factors such as natural diet, feeding rates, reproduction, and growth are well understood.     

Testes mass in megachiropteran bats varies in accordance with sperm competition theory

Sperm competition is a widespread phenomenon influencing a range of characters, including investment in gonadal tissue. Conspecific proximity is one factor which can influence the risk of sperm competition and hence testicular investment, and decreased confidence of paternity may be one cost of group living. Aspects of female biology may also influence spermatogenic investment and sperm morphology. This study examines the associations between relative testes mass and roost-group size across 17 species of Megachiroptera. Associations between breeding season duration and investment in spermatogenesis are also examined, as are associations between female reproductive tract dimensions and testes mass and dimensions of spermatozoa across all bats. Relative testes mass was significantly positively associated with roost-group size at a species level and after appropriate phylogenetic control (pairwise comparisons and comparison of independent contrasts). There were no significant relationships between breeding season duration and relative testes mass. Across all bats, neither testes mass nor sperm length were significantly related to dimensions of the female tract. The results are discussed in the context of sperm competition. Received: 7 January 1998 / Accepted after revision: 8 August 1998     

Effects of temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis

To assess the effects of both temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis (O.F. Müller), individuals collected December 1996 (winter experiment) and June 1997 (summer experiment) were maintained for 3 months in one of four experimental treatments: (1) 3 °C and fed ad libitum (high ration), (2) 3 °C and fed one-seventh of the maximum ration (low ration), and (3) 12 °C and fed the high ration; (4) 12 °C and fed the low ration. All individuals were fed an artificial diet and exposed to only 1 h of light every day. At the end of both experiments, mean gonad indices of sea urchins fed the high ration had increased significantly (11–24% and 6–19% in the winter and summer experiments, respectively), while the gonad indices of individuals fed the low ration did not change. At the high ration (both experiments), the increase in gonad index of sea urchins occurred primarily as the result of a significant increase in the mass of nutritive phagocytes, as revealed by histological analyses. Primary oocytes were significantly larger in individuals held at 3 °C than at 12 °C throughout the winter experiment, regardless of food ration; during the summer experiment, primary oocytes were significantly larger in individuals receiving the high ration, regardless of the temperature at which they were held. These results suggest that: (1) food availability is the most important factor regulating energy storage and the relative size of gonads throughout the year, (2) temperature affects the rate of growth and maturation of primary oocytes during the later stages of oogenesis, and (3) once gametogenesis has been initiated, mature ova will be produced, even under conditions of low food availability. Conditions of high food availability in summer and low temperature in winter would thus favor reproductive output in sea urchin populations. Received: 1 March 2000 / Accepted: 4 October 2000     

Effects of inorganic and organic phosphates on feeding, feeding absorption, nutrient allocation, growth and righting responses of the sea urchin Lytechinus variegatus

The sea urchin Lytechinus variegatus is capable of surviving chronic exposure to sodium phosphate (inorganic phosphate) concentrations as high as 3.2 mg l⁻¹, and triethyl phosphate (organic phosphate) concentrations of 1,000 mg l⁻¹. However, chronic exposure to low (0.8 mg l⁻¹ inorganic and 10 mg l⁻¹ organic phosphate), medium (1.6 mg l⁻¹ inorganic and 100 mg l⁻¹ organic phosphate) or high (3.2 mg l⁻¹ inorganic and 1,000 mg l⁻¹ organic phosphate) sublethal concentrations of these phosphates inhibits feeding, fecal production, nutrient absorption and allocation, growth and righting behavior. Food consumption and fecal production declined significantly in individuals exposed to medium and high concentrations of inorganic phosphates and all levels of organic phosphates. Feeding absorption efficiencies for total organics and carbohydrates decreased significantly in individuals held in the highest concentration of organic phosphate. Feeding absorption efficiencies for lipids were significantly reduced in the highest inorganic phosphate concentration only, while they decreased significantly for protein with increasing phosphate exposure. Carbohydrate and lipid levels in gonad and gut tissues decreased significantly with exposure to increasing phosphate concentrations, potentially impairing both gametogenesis and nutrient storage in the gut. Moreover, gonad indices significantly decreased in individuals exposed to the highest concentrations of either phosphate. Growth rates decreased significantly under the influence of all phosphate concentrations, while increasing in seawater alone. Individuals exposed to increasing phosphate concentrations showed reduced righting responses (a measure of stress) and no acclimation in righting times during chronic exposure to phosphates over a 4 week period. These findings indicate that shallow-water populations of L. variegatus subjected to inorganic and organic phosphate pollutants will exhibit stress and be inhibited in their growth and performance due to reductions in feeding, nutrient absorption and allocation of nutrients to key somatic and reproductive tissues. Received: 10 April 2000 / Accepted: 2 October 2000     

Reproductive biology of the edible jellyfish Catostylus mosaicus (Rhizostomeae)

 The reproductive biology of the jellyfish Catostylus mosaicus (Quoy and Gaimard 1824) (Scyphozoa: Rhizostomeae) was investigated in New South Wales, Australia. Medusae were gonochoristic. There was a 1:1 ratio of male and female medusae and there was no evidence of sexual dimorphism. Oocytes arose from the gastrodermis and maintained contact with the gastrodermis, via trophocytes, throughout gametogenesis. Spermatogenesis occurred within follicles that arose from invagination of the gastrodermis. Detailed sampling of gonads over a period of 3.25 yr in Botany Bay, and over 2.5 yr in Lake Illawarra, indicated that gametogenesis occurred almost continuously during the year. Oocytes were smaller, or were absent from the ovaries during 3 of the 4 winters sampled at Botany Bay and during all 3 winter periods sampled at Lake Illawarra. Comparisons were made with other locations, although these were sampled less frequently. When medusae were present at a location, similar trends were observed. The size at which medusae matured varied, but during non-winter periods and at two locations, all medusae exceeding 130 mm diam were considered mature. Received: 6 January 2000 / Accepted: 3 July 2000     

Importance of cysts in the population dynamics of the red tide flagellate Heterosigma akashiwo (Raphidophyceae)

To elucidate roles of cysts in occurrences of Heterosigma akashiwo blooms, cyst dynamics were studied in northern Hiroshima Bay, the Seto Inland Sea of Japan, where H. akashiwo regularly forms red tide in June. Monthly measurements of seasonal changes in the densities of vegetative cells of H. akashiwo and their germinable cysts in surface sediments (top 1-cm layer) were made for 2 years at three stations. Vegetative cells of H. akashiwo could be detected from April through December throughout the water column, and the existence of vegetative cells was confirmed in surface waters even in winter after incubation of sampled seawater in culture medium. Germinable cysts, enumerated by the extinction dilution method, existed in sediments in all seasons, even before and after the seasonal bloom. The effects of incubation temperature on the germination of natural cysts of H. akashiwo in sediments were examined. Germination was not observed at 5 °C, was low at 10 °C, while it increased at 15 °C, and maintained a high level to 25 °C. The bottom water temperature reached 15 °C (suitable for the germination of cysts) and the surface about 18 °C or more (suitable for the growth of vegetative cells) 2 to 3 weeks before the blooms. The dark survival of H. akashiwo cysts was tested, and it was found that the cysts were viable for at least 650 d at 11 °C, and for 165 d at 25 °C, indicating a significant role of cysts in the survival during winter and summer seasons. The cysts presumably also play an important role in seeding primary populations into water columns when the bottom water reaches a suitable temperature (around 15 °C); thereafter the populations develop with great annual regularity to bloom in June. These results suggest that initiation of H. akashiwo red tides in the Seto Inland Sea could be triggered by bottom water temperature. Received: 3 July 1998 / Accepted: 12 January 1999     

Effects of sudden changes in salinity on endogenous rhythms of the spotted sea bass Lateolabrax sp.

The endogenous rhythm of oxygen consumption in juvenile spotted sea bass (Lateolabrax sp.) was measured to test the effects of sudden changes in salinity on the metabolic activity. Mean oxygen consumption rates of this euryhaline fish decreased by 13.5 to 16.0% and 25.3 to 36.4% when they were transferred from 31.5 to 15‰ seawater and to fresh water (0‰), respectively. The maximum rate of oxygen consumption was observed between 18:00 and 19:00 hrs local time, 1 to 2 h before sunset, even though they were kept in constant darkness. The peaks of oxygen consumption occurred in 23.2- and 23.3-h intervals, which correspond with a circadian rhythm, as revealed by maximum entropy spectral analysis. A markedly weakened rhythm in oxygen consumption occurred from 8 to 10 d after onset of the experiments. This study indicates that spotted sea bass can withstand sudden drops in salinity from 31.5‰ to fresh water, and yet maintain a regular though somewhat dampened endogenous rhythm of oxygen consumption. Received: 16 June 1997 / Accepted: 3 February 1998     

Diet and egg production of the copepod Acartia tonsa in Florida Bay. II. Role of the nutritional environment

 As part of an ongoing study of changes in the trophic pathways of Florida Bay's pelagic ecosystem, the nutritional environment (seston protein, lipid and carbohydrate levels), diet (taxon-specific microplankton ingestion rates) and egg production rate of the important planktonic copepod Acartia tonsa were measured off Rankin and Duck Keys in July and September 1997 and in January, March and May 1998. Rankin Key has been the site of extensive sea grass mortality and persistent ultraplankton blooms since 1987. Duck Key has experienced neither of these perturbations. Protist (auto-plus heterotroph) biomass was approximately twice as high off Rankin as off Duck Key. Diatoms, dinoflagellates and heterotrophic protists dominated the food environment off Rankin Key, while cells <5 μm diam often predominated off Duck Key. Protein and carbohydrate concentrations were higher off Rankin Key than Duck Key, while average lipid levels were usually low at both stations. Ingestion rates at both stations frequently approached temperature- and food-dependent maxima for the species, exceeding 100% of estimated body C d⁻¹ on 3 of 5 occasions off Rankin Key. Egg production rates, however, were consistently low (Rankin: 3 to 16 eggs copepod⁻¹ d⁻¹; Duck: 1 to 12 eggs copepod⁻¹ d⁻¹), and gross egg production efficiencies (100% × egg production C/ingested C) averaged <10%. At Duck Key, egg production rate varied with temperature and food concentration, while off Rankin Key, egg production was strongly correlated with seston protein content. The efficiency with which lipids (which were scarce in the seston) were transferred from the diet to the eggs increased exponentially with decreasing seston lipid content. Egg production efficiencies based on protein, however, were independent of seston protein content and never exceeded 10%. Received: 23 December 1998 / Accepted: 23 March 2000     

Reproduction and growth of Isostichopus fuscus (Echinodermata: Holothuroidea) in the southern Gulf of California, México

The reproduction and growth of Isostichopus fuscus (Ludwig, 1875) at Isla Espíritu Santo, Baja California Sur, México from July 1992 to September 1993 and from August 1996 to July 1997 were analyzed. The reproductive cycle was monitored by using histological analysis. Five gonadal stages were described according to cell types present in the gonad; undetermined, gametogenesis, maturity, spawning and post-spawning. This holothurian reproduces annually during summer (July to September), when sea surface temperature reaches 27 °C. The gonad index is related to the gonadal stages. The first sexual maturity for I. fuscus was calculated at 367 g (4 to 5 years old) according to the von Bertalanffy equation, and nearly all the specimens at Isla Espíritu Santo were 5 to 7 years old (including some individuals over 35 years old). Data of the length–weight relationship showed that I. fuscus grows allometrically at the studied site (b = 1.83). The mortality rate was relatively high (M = 0.51), indicating that in a typical cohort, 40% of the recruits does not survive the first year, and less than 20% reaches maturity. Received: 15 July 1998 / Accepted: 19 August 1999