Ammonia as confounding factor in toxicity tests with the sea urchin Paracentrotus lividus (Lmk)

The aim of this study was to investigate the toxicity of total ammonia towards sea urchin bioassays, in order to elucidate the role of ammonia as confounding factor in sediment quality assessment studies. New toxicity data, expressed as EC50, NOEC and LOEC are reported for Paracentrotus lividus based on experiments at differing pH values. Results confirmed that sperm cell toxicity test is not very sensitive to ammonia, while total ammonia embryotoxicity is strictly pH dependent. Accurate observations of embryotoxic effects at increasing total ammonia concentrations evidenced progressive shifts from malformed plutei to gastrula and blastula blockages. Toxicity data obtained for P. lividus are comparable with previous results with other echinoid species.     

Fish predators and scavengers of the sea urchin Paracentrotus lividus in protected areas of the north-west Mediterranean Sea

Direct observations of predation on 436 individuals of the sea urchin Paracentrotus lividus (Lamarck) were carried out in infralittoral rocky bottoms (between 5 and 20 m deep) in three Mediterranean marine reserves. The predator guild was composed of six fish species, the sparids Diplodus sargus and D. vulgaris being the main predators, and the labrid Coris julis a major predator of juvenile sea urchins. Four species attempted but failed to open sea urchins. The scavenger guild was most rich in species, with 17 species observed. Predation was size-dependent; the size of predators increased with increasing size of the sea urchins. The presence of two feeding guilds is suggested, one composed of sparids (Diplodus spp.), able to kill juvenile and adult sea urchins, and the other composed of labrids (mainly C. julis), which feed on juvenile sea urchins. To avoid the extension of overgrazed, barren areas created by P. lividus populations, fisheries' regulations should focus on major sea-urchin predators, chiefly D. sargus, D. vulgaris and C. julis. Received: 23 April 1997 / Accepted: 30 May 1997     

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     

Reproductive dynamics of the sea urchin Paracentrotus lividus on the Galicia coast (NW Spain): effects of habitat and population density

We studied the spatial variability in the size at first maturity and the reproductive cycle of Paracentrotus lividus in Galicia, contributing key information for the exploitation and management of this resource. The size at maturity varied between 20.4 (±1.2 SE) mm and 27.9 ± 1.2 mm and was smaller in areas of low population density where sea urchins do not form patches. Using a nonlinear model, we analysed the effect of depth, body size, sex and population density on the temporal pattern of the gonad index. The maximum and minimum indices were obtained at 4 m depth in the months before and after the spring spawning, respectively. The depth also affected the cycle phase, and the sea urchins at 4 m depth spawned 9.4 ± 3.0 days later than the sea urchins at 8 m depth and 20.5 ± 3.0 days later than those at 12 m depth. Moreover, the sea urchins living in patches showed a slight increase in gonad size as a consequence of the better-quality habitat. This shows that there is no intraspecific competition in this area despite the high population densities reached (18.5 kg m^?2).     

Temperature, salinity and growth rate dependences of Mg/Ca and Sr/Ca ratios of the skeleton of the sea urchin Paracentrotus lividus (Lamarck): an experimental approach

The skeletal Mg/Ca ratio of echinoderms is known to increase with temperature but the relation has never been established in controlled experimental conditions. The present study investigated the effect of temperature, salinity and growth rate on Mg/Ca and Sr/Ca ratios in calcite skeletons of juvenile sea urchins grown in experimental conditions. Mg/Ca ratio was positively related to temperature, increasing until a plateau at high but field occurring temperatures. It was not linked to salinity nor growth rate. We suggest that this plateau is due to properties of the organic matrix of mineralization and recommend to take it into account for the use of Mg/Ca as proxy of seawater Mg/Ca. Skeletal Sr/Ca ratio was mainly dependent on temperature and growth rate, as usually observed in calcite skeletons.     

Profitability and sustainability of edible sea urchin fishery in Sardinia (Italy)

Sea urchin (Paracentrotus lividus) fishery is intensively practiced in several areas of the Mediterranean basin. In Sardinia, as well as other Mediterranean countries, sea urchin is a basic ingredient for several dishes due to the delicacy of its gonads (roe), and demand is constantly increasing. Restrictions have been implemented in order to minimise the risk of overexploitation, however, these measures might jeopardize economic convenience in sea urchin harvesting. This paper estimates economic convenience within the edible sea urchin fishery in Sardinia. It aims to determine whether both profitability and sustainability, in terms of the preservation of sea urchin stock, can be guaranteed by actual policy regulation. We found high variability in terms of captures and profitability among firms, and a considerable degree of this variability is the result of technological differences between fishing methods. Analysis also suggests that a slight increase in allowed captures should generate a more than proportional increase in profits. This evidence gives some useful suggestions for improving the efficacy of policies in affecting the economic and environmental sustainability of the Mediterranean sea urchin fishery.     

Evidence of antibacterial and lysozyme-like activity in different planktonic larval stages of Paracentrotus lividus

Larval lysate (four-armed stage) exerts a lysozyme-like and antibacterial activity directed against Vibrio alginolyticus and other bacterial strains. This lysozyme activity is dependent on the pH and ionic strength of the reacting medium and sample. The other larval stages (six-armed, eight-armed and a stage approaching metamorphosis) possess protection against bacteria as well. The highest antibacterial activity is present in the six-armed stage. These antibacterial compounds avoid exploitation by bacteria. Paracentrotus lividus larvae used in our experiments were laboratory-reared from specimens collected in 1993 in Porto Cesareo (Lecce, Italy).     

CO2 leakage simulation: effects of the pH decrease on fertilisation and larval development of Paracentrotus lividus and sediment metals toxicity

Carbon capture and storage has become a new mitigation option to reduce anthropogenic CO₂ emissions. The effects of the CO₂-related acidification, associated with unpredictable leaks of CO₂ during the operational phases were studied using the Paracentrotus lividus sea urchin-liquid-phase-assays (fertilisation and embryo-larval development tests). Besides to the direct effects of the CO₂-associated pH decrease, the possible effects on marine sediment toxicity were studied by exposing the urchin larvae to elutriate of sediments with different metals concentration, which were subjected to various pH treatments. The resulted median toxic effect pH ranged from 6.33?±?0.02 and 6.91?±?0.01 for the egg fertilisation, and between 6.66?±?0.03 and 7.16?±?0.01 for the larval development assays. Concentrations of Co, Zn, As, Cu, and Fe were detected in the sediment elutriates. For all the sediment elutriates tested the amount of the dissolved Zn increased in parallel with the pH reductions. Correlated effects of acidification, biological response, and the presence of dissolved metals were observed in this work evidencing that the most important factor controlling the reduction of egg fertilisation and larval development success was the combined effects of the reduction of pH and the presence of the contaminants (mainly Zn, Co, and As).     

Geographic,population, and seasonal patterns in the reproductive parameters of the sea urchin <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>

Generalized additive models were used to synthesize the data from 52 publications on the gonad growth and spawning seasonality of Paracentrotus lividus and identify spatial and temporal patterns in the reproductive processes. According to our results, Atlantic populations spawn in spring and develop large gonads (6.4 ± 0.2% of fresh body weight (mean ± SE)), which increase in size toward higher latitudes. While in the Mediterranean, the gonads are smaller (3.1 ± 0.1%), possibly as the result of successive spawnings which impede nutrients accumulating in the gonads. Differences were also observed between habitats: gonad production was higher in seagrass meadows in the Mediterranean and in subtidal rocky substrates in the Atlantic. Food availability might cause these variations as well as the negative correlation observed between depth and the gonad index for rocky substrates. Sex and body size do not seem to influence greatly gonad growth.     

Patterns of abundance, population size structure and microhabitat usage of Paracentrotus lividus (Echinodermata: Echinoidea) in SW Portugal and NW Italy

In coastal habitats, wave exposure influences several aspects of the life history of marine organisms. Here, we assess how hydrodynamic conditions can generate variation in density, size structure and microhabitat usage of Paracentrotus lividus and whether these effects are consistent between regions that are markedly different for oceanic climate, such as the coasts of SW Portugal and NW Italy. The abundance of P. lividus was ~4 times higher in SW Portugal than in NW Italy, but within each region, there was no effect of wave exposure. In SW Portugal, higher urchin abundances were found at shallower depths, while no effect of depth on urchin abundance emerged in NW Italy. Most of the variation in urchin abundance occurred at small spatial scales (metres), and our results suggest that habitat complexity, that is, the presence of cracks and crevices, is an important determinant of patterns of distribution of this species. The population in NW Italy was characterized by a unimodal size distribution, with a higher proportion of medium-sized individuals. In contrast, in SW Portugal, smaller individuals represented a large proportion of the populations. Size structure varied between exposed and sheltered habitats in SW Portugal, suggesting that the proportion of individuals from different size cohorts may vary along wave-exposure gradients as a result of direct or indirect effects of hydrodynamic forces. In SW Portugal, most urchins occurred in burrows, while in NW Italy, urchins were mainly observed in crevices. These results suggest that creating/occupying burrows might be an adaptive behaviour that allows sea urchins to better withstand stressful hydrodynamic conditions and, therefore, are more common on exposed Atlantic coasts. Overall, our study suggests that the effects of hydrodynamic forces on sea urchin populations are context dependent and vary according to background oceanic climate.     

Circulation of perivisceral fluid in the sea urchin Lytechinus variegatus

The principal driving mechanism of perivisceral-fluid circulation was investigated in the echinoid Lytechinus variegatus by metal-clad hot-wire anemometry. Analysis of 23 flow profiles from 13 individuals collected from 1 to 3 m water depth at Tarpon Springs and Sarasota, Florida (USA) in 1984 indicates that for this species, the ciliated epithelium is an ineffective driving mechanism for mixing of the perivisceral fluid. Instead, the Aristotle's lantern is the major driving force of circulation of the perivisceral fluid, with velocity fluctuations of 1.5 mm s^-1 amplitude in the cavity center during intermittent periods of activity. These fluctuations rapidly decay as the ciliated surfaces are approached. Spectral analysis of velocity time-series reveals predominant energy-containing velocity fluctuations between 0.007 and 0.05 Hz. The circulation, which can be described as non-turbulent forced mixing, has revealed four different flow patterns which appear to be correlated with specific animal activities. A modified eddydiffusion model predicts that oxygen transport through the cavity over a 2 cm path length requires more than 20 h. Calculation of the oxygen flux to a ripe ovary suggests that supply does not meet ovarian oxygen demand in this case. Alternative pathways of oxygen transport invoking advection in perivisceral-fluid eddies and along the ciliated coelomic surfaces are compatible with the observed flow patterns and lead to much shorter oxygen-transport time scales of the order of 1 to 10 min.     

Experimental evaluation of aggregation behavior in the sea urchin Strongylocentrotus droebachiensis

Defensive aggregation by Strongylocentrotus droebachiensis has been invoked as the fulcrum for the transformation of macroalgal beds into coralline barrens in the northwest Atlantic. We critically examined some of the mechanisms contributing to aggregation behavior by experimentally manipulating sea urchins, purported predators and food during autumn 1983 and spring 1984 both in the laboratory and field. We utilized several approaches to examine a range of sea urchin responses to the presence of food, tethered predators, caged predators, crushed con-specifics in the field and predators in laboratory tanks. Some of the field cages had the property of allowing free passage to sea urchins while retaining lobsters; this allowed distinctions to be made between artifacts caused by cage walls or topographic barriers and unrestricted behavioral responses of sea urchins. The results falsified the hypothesis that aggregations of S. droebachiensis are elicited by predators. Except in the presence of algae, sea urchins always avoided decapod predators (but not sea stars) and fled from the vicinity of injured conspecifics. However, avoidance behavior was subordinate to feeding responses, demonstrated by mass migrations of sea urchins into cages (with lobsters) to feed on algae. We reaffirmed by observation and manipulation previous studies which showed that sea urchins aggregate only in the presence of food. Two types of sea urchin groupings were delineated: (1) surficial two dimensional associations, often caused by topographic or other features which inhibited dispersal and (2) cohesive three-dimensional aggregations induced by food.     

Changes in sea urchin populations after the destruction of kelp beds

In St. Margaret's Bay, Nova Scotia, Canada, there are large areas in which sea urchins (Strongylocentrotus droebachiensis) have eliminated beds of kelp (Laminaria spp.). Sites were identified where destruction of kelp beds had taken place 1, 2, 3, 3.5 and 4 years ago. With increase of time since kelp disappearance, the sea urchins showed decreased growth rate, reduced gonad size, but an increase in numbers resulting from high recruitment rates in the first two years after kelp bed destruction. These sea urchin populations, by their browsing, effectively prevent the regeneration of kelp. There is, as yet, no evidence of the sea urchin populations being starved out to allow the kelp to return.     

Reproductive cycle of the sea urchin Holopneustes purpurascens (Temnopleuridae: Echinodermata)

This study documents the annual reproductive cycle of the Temnopleurid echinoid Holopneustes purpurascens near Sydney. H. purpurascens reached sexual maturity between 11 and 26 mm diameter and was gonochoristic. The sequence of oogenesis was similar to previously described echinoids. However, the proportion of eggs at each stage showed no temporal pattern in abundance, suggesting that vitellogenesis is continuous throughout the year. Spermatogenesis was also similar to other echinoids and showed a significant increase in the proportion of partially spawned testes in spring, followed by a similar increase in the proportion of partially spawned testes in late spring and summer. Gonad indices showed a peak in weight from late winter to mid-spring (9.10ǃ.47 for females; 6.28ǂ.43 for males; mean-SE), then a decline for both sexes. Although spawning was variable over time, there was a peak in the proportion of spawning from late winter to early summer that correlated with the observed peaks in gonad indices and gametogenic cycles. The demographic implications of this reproductive pattern are that new recruits can enter the population through synchronous spawning during the breeding period, and smaller haphazard spawning events at other times of the year.     

Foods and predators of the green sea urchin Strongylocentrotus droebachiensis in Newfoundland waters

Gut analyses of the green sea urchin Strongylocentrotus droebachiensis (O. F. Müller) demonstrated that perennial phaeophytes, mostly fucoids and Alaria esculenta, were predominant in the diet. Ephemeral species, coralline algae and animals, were consumed in smaller amounts when available. Grazing by the urchins is evidently responsible for the dearth of non-coralline sublittoral algae in Newfoundland waters. Lobsters, rock crabs, purple sea stars, other urchins, and a variety of fishes and birds feed on S. droebachiensis, but predation is apparently not effective in limiting the abundance of the urchin.Studies in Biology from the Memorial University of Newfoundland No. 234.Contribution from the Marine Sciences Research Laboratory No. 66.     

Anaerobic and aerobic energy metabolism in ovaries of the sea urchin Strongylocentrotus droebachiensis

The gonads of sea urchins undergo large changes in mass during their gametogenic cycle. In addition, they have relatively low aerobic capacities and are poorly perfused by the circulatory system and thus are continually hypoxic or anoxic. The present study of Strongylocentrotus droebachiensis investigates seasonal changes in the relationships among mass of the ovaries, pH and P_O2 of the perivisceral coelomic fluid which bathes the ovaries, and partitioning of ovary energy metabolism into its anaerobic and aerobic components. S. droebachiensis were collected at Blue Hill Falls, Maine, USA, from August 1982 to March 1984. We found that from 76 to 92% of the heat dissipated by isolated ovaries of the sea urchin S. droebachiensis derives from anaerobic energy metabolism at partial pressures of oxygen prevailing in vivo. Ovaries from S. droebachiensis have the capacity to produce large amounts of lactate under imposed anoxia, but lactate accounts for only 37% of the total anoxic heat dissipation, which suggests that other end products of anaerobiosis are present. Seasonal changes in pH and P_O2 of the perivisceral coelomic fluid can be explained by a complex relationship among changes in temperature, reproductive condition, and anaerobic and aerobic metabolism in the ovaries, gut and body wall. Seasonal changes in the buffering capacity of the perivisceral coelomic fluid must be determined before the effects of respiratory and metabolic acid production on the acid-base status of the coelomic fluid can be fully understood.     

Resource allocation and growth rates in the sea urchin Evechinus chloroticus (Echinoidea: Echinometridae)

The morphometry of the sea urchin Evechinus chloroticus from habitats of contrasting algal abundance but of similar urchin density was examined at two localities in southern New Zealand during 1993. Populations from habitats of high algal abundance (Dusky Sound) had similar relationships of demipyramid (jaws) to test diameter, but individuals had significantly smaller jaws relative to their test diameter than those from a locality where algal abundance was low (Arapawa Island). The body wall mass (in relation to total wet weight) was similar for populations from both localities but, for Dusky Sound populations, individuals from exposed sites had greater relative body mass than those from sheltered sites. The ratios of gonad weight:total weight were similar between localities. However, E. chloroticus from Arapawa Island reached reproductive maturity at a smaller size than those from Dusky Sound. Growth rates of E. chloroticus varied among localities in Dusky Sound. Growth was modelled by the Tanaka function, which allows for rapid early growth until reproductive maturity is reached and declining growth rates thereafter. The results show that sea urchins respond to low resource availability by increasing the size of the food-gathering apparatus, maturing at a smaller size, and growing to a smaller size than individuals from food-rich habitats. Received: 3 December 1996 / Accepted: 29 January 1997     

Karyotype analysis of the sea urchinParacentrotus lividus (Echinodermata): evidence for a heteromorphic chromosome sex mechanism

A consistent diploid number of 2n = 36 was determined for the sea urchinParacentrotus lividus from the Gulf of Palermo by analysis of mitotic chromosomes of both early developing embryos and male gonads. The haploid numbern = 18 was determined by counts of spermatocyte bivalents at diakinesis. A heteromorphic chromosome sex mechanism of the XY type is likely present in this species. This is indicated by the occurrence of a chromosomal pair, pair No. 2, which is heteromorphic in both morphology and size in about 50% of the mitotic figures (metaphases and anaphases) of einbryos. In addition, heteromorphism of the same pair of chromosomes occurred during spermatogonial metaphases in the five male specimens investigated here. The detection of a low chromosome number (2n = 36) compared to other echinoids (2n = 42 to 44), a heteromorphic chromosome sex system and the involvement of three chromosome pairs in nucleolar organization (NORs) provide evidence of the specialization of theP. lividus karyotype.     

Skeletal calcification patterns in the sea urchin Tripneustes gratilla elatensis (Echinoidea: Regularia)

A comprehensive study on skeletal calcification in the regular echinoid Tripneustes gratilla elatensis was carried out between 1979 and 1981 in the northern Gulf of Eilat (Red Sea), employing size measurement, allometry and radiotracer techniques. Uptake rates of the isotope ⁴⁵Ca were used to estimate the calcification rate of whole tests. Calcification rates of the different skeletal parts and the ⁴⁵Ca uptake along the sutural margins of individual plates were also measured. Whole-test calcification rates for juvenile individuals, 11 to 13 mm in diameter, relative to their skeletal Ca dry weight, calculated using the allometric relationship between skeletal dry weight and diameter, were 1.5 to 1.8% d^-1 for 4 to 8 h of incubation, while calcification rates obtained from periodical size measurements and allometrical constants amounted to 2.75% d^-1. The apparent discrepancy between these results is explained partly by the short duration of the incubations, during which the internal calcium pools, apparently in the coelom, were not fully saturated with the radioactive tracer. This discrepancy decreased when longer radioactive incubations were used, or when a post-incubational deposition of ⁴⁵Ca from these pools was allowed. The size of these pools is roughly 10 mol Ca per 13-mm individual, and it resides probably in the coelom where Ca concentration is 3 times higher than in seawater. The calcification rates given as percentage of skeleton added per day, both for the radioactive and allometric methods, decreased with the urchins' size. The calcification patterns of various components of the skeleton generally fitted their allometrical trends — a relative decrease in the size of the spines, apical plates and lantern. The teeth calcify rapidly to compensate for the constant erosion of their tips. Calcification per plate decreased exponentially from the apex to the peristome. Plate-edge calcification patterns of both interambulacral and ambulacral plates fitted patterns derived from size increment measurements, using natural growth-lines. Typically, most plates grow more horizontally (latitudinally) than vertically. It is suggested that the vertical vs horizontal calcification ratio (v/h) determines the individual plate growth and affects the whole test morphology.