Oxygen and acid-base status of the sea urchin Psammechinus miliaris during environmental hypoxia

The respiratory and acid-base responses to hypoxia of the sea urchin Psammechinus miliaris Gmelin have been studied both in the laboratory and in the field. Sea urchins were collected from the Clyde Sea area, Scotland, between March and July 1993. Individual urchins were unable to regulate their oxygen uptake during hypoxia. The partial pressure of oxygen (P_{O 2}) in the coelomic fluid was lower than ambient P_{O 2} during normoxia but still decreased with a decrease in environmental P_{O 2}. There was a significant increase in pH and in the concentrations of bicarbonate, magnesium and calcium in the coelomic fluid, but a significant decrease in the partial pressure of carbon dioxide (P_{CO 2}), in response to declining P_{O 2}s. The disturbance of respiratory parameters in urchins accompanying hypoxic exposure in rock pools was similar to that observed in the laboratory, although not as severe as might have been expected on the basis of the laboratory experiments.     

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.     

Skeletal changes in the test and jaws of the sea urchinDiadema antillarum in response to food limitation

When food-limited,Diadema antillarum increases the relative size of Aristotle's lantern compared to the size of the test. This is accomplished primarily by decreasing the size of the test: the demipyramid grows at a reduced rate as the test shrinks. These results suggest that an altered relationship between demipyramid and test size can provide evidence for food-limitation in field populations. The sizes of the lantern and test of museum and fossil specimens can be investigated for the presence of food limitation and biotic interactions in recent and past times.     

Skeletal growth markers in the deep-sea brittle starsOphiura ljungmani andOphiomusium lymani

Growth bands have been found in the calcitic vertebral arm ossicles of the commonly occurring deep-sea brittle starsOphiura ljungmani Wyville Thomson andOphiomusium lymani (Lyman) (Echinodermata: Ophiuroidea) trawled from the Rockall Trough (N.E. Atlantic) at 2 200 and 2 900 m depth from 1973 to 1982. InOphiura ljungmani, the study of ossicle microstructure by SEM shows that growth bands reflect differences in stereom porosity and surface relief, similar to that previously found amongst shallow-water brittle stars. The pattern inOphiomusium lymani was much less clear from the microstructure, but could be revealed by heating the ossicle to 450 °C and clearing in xylene. The bands showed up as fine, translucent rings separated by more opaque, wider zones, perhaps reflecting differences in organic material incorporated within the calcite. Both growth-banding patterns probably reflect an annual cycle in skeletal growth rate. On this assumption, the number and spacing of the banding in the two species indicates contrasting growth strategies. WithOphiura ljungmani, a rather regular annual growth increment and perhaps shorter lifespan (up to ca. 10 yr) thanOphiomusium lymani is indicated. The latter shows a relatively wide spacing of early bands, followed by tight clustering of the outermost bands corresponding to adult sizes. This growth pattern is characteristic of species escaping from predation by rapid growth to relatively large adult size. AdultO. lymani probably grow slowly, some perhaps reaching 20 yr of age. Growth curves were fitted to size-at-age corresponding to measurements of the size and ordering of growth bands. These corroborate age structure previously estimated from analysis of size frequencies in time series from the stations sampled in the present study. Such skeletal growth markers should be of value in analysis of the demography of deep-sea populations.     

RNA-DNA ratio: an index of larval fish growth in the sea

Data on water temperature, RNA-DNA ratio, and growth of eight species of temperate marine fish larvae reared in the laboratory were fit to the equation: $$G_{pi} = 0.93{\text{ }}\operatorname{T} + 4.75{\text{ RNA - DNA}} - 18.18$$ where G_pi is the protein growth rate in % d^-1 and T is the water temperature. Water temperature and larval RNA-DNA ratio explained 92% of the variability in growth rate of laboratory-reared larvae. The model is useful over the entire range of feeding levels (starvation to excess), temperatures (2° to 20°C) and fish species studied. Estimates of recent growth of larval cod, haddock, and sand lance caught at sea based on water temperature and RNA-DNA ratio ranged from negative to 26% d^-1. These data demonstrate the importance of food availability in larval fish mortality and suggest that short-term growth under favorable conditions may be considerably higher than expected from long-term indicators. RNA-DNA ratio analysis offers new possibilities for understanding larval growth and mortality, and their relation to environmental variability.     

Natural growth bands and growth variability in the sea urchin Echinus esculentus: results from tetracycline tagging

Growth of the European edible sea urchin Echinus esculentus L. was studied in a population held for 2 yr in cages on the sea bed, after labelling with the skeletal growth marker tetracycline. The final position of the tetracycline tag on the genital plates agreed with an annual periodicity in natural growth zones; two such growth zones appeared beyond the position of the tag on the ground surface of the plate as light-reflecting bands separated by narrow dark lines in the middle layer. Individual and group (pooled data) growth parameters were estimated from the growth increment shown in the genital plate, whose lateral growth displayed a linear relationship to the diameter of the urchin test within the size range of these measurements. Von Bertalanffy growth parameters (asymptotic size and growth-rate function K) fitted to the growth increment on each individual were highly significantly correlated to those fitted to the natural growth lines, assuming an annual periodicity. The inferred growth pattern agrees well with conclusions based on H. B. Moore's growth-band data. The caged urchins can be assumed to have experienced exactly similar conditions, yet the growth curves fitted to individuals showed considerable variability. The good agreement between estimated growth function parameters of individuals obtained by the two methods indicate that this reflects real variability in growth between individuals that probably is genetically rather than environmentally determined. The growth of E. esculentus, and the adaptational significance of high growth variability in the population is briefly discussed.     

Photoperiodic regulation of gametogenesis and gonadal growth in the sea star Pisaster ochraceus

A photoperiod regime 6 mo out of phase between December 1978 and August 1980 resulted in gametogenesis, gonadal growth, and spawning in laboratory-maintained sea stars [Pisaster ochraceus (Brandt)] 6 mo out of phase with individuals in the field or laboratory on normal celestial photoperiods at Santa Cruz, California (USA). The seasonal fluctuation of pyloric cecum size also was shifted when the sea stars were held on the out-of-phase photoperiod regime. Phase shift of all these events was evident within 6 to 9 mo. Long daylengths (or short nightlengths) in spring and summer apparently synchronize or entrain the initiaion of gametogenesis and gonadal growth in fall.     

Effects of diet on fatty acid composition of body zones in larvae of the sea bass Dicentrarchus labrax: a chemometric study

Larvae of the sea bass Dicentrachus labrax were fed four Artemia sp. diets for 28 d. Three were nauplii enriched with emulsions of polyunsaturated fatty acids, and the fourth nauplii enriched with baker's yeast. At the end of the experimental period, the fatty acids of the bodies, heads and eyes of the larvae were analysed. A multivariate statistical method (discriminant analysis, DA) applied to the data revealed anatomical as well as dietary fatty acid pattern-discrimination. We propose here the use of discriminant analysis as a pattern-recognition method that will help to integrate the fatty acid information obtained in nutritional studies.     

Morphological adaptation of a planktonic diatom to growth in Antarctic sea ice

Chaetoceros dichaeta Ehrenberg is one of the most important planktonic diatom species in the Southern Ocean, making a significant contribution to the total biomass in the region. Our observations on both field and culture material have revealed the existence of a specialized form of C. dichaeta adapted to living in sea ice. This sea ice form differs from the planktonic form by the shape and orientation of the setae and the aperture length between sibling cells. Thus, the diameter of the chain is equivalent to the apical axes of the cells and is accompanied by a two order of magnitude decrease in minimal space requirement. Here, we report for the first time on the extraordinary overwintering strategy of a planktonic diatom in sea ice facilitated by its rapid morphological adaptation to changing environmental conditions. This morphological plasticity enables it to thrive in the confined space of the sea ice brine matrix and retain its numerical dominance in recurrent growing seasons and has likely evolved to optimally exploit the dynamic ecosystem of the seasonally ice-covered seas of the Southern Ocean.     

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     

Modelling the effect of fibropapilloma disease on the somatic growth dynamics of Hawaiian green sea turtles

The effect of the tumour-forming disease, fibropapillomatosis, on the somatic growth dynamics of green turtles resident in the Pala’au foraging grounds (Moloka’i, Hawai’i) was evaluated using a Bayesian generalised additive mixed modelling approach. This regression model enabled us to account for fixed effects (fibropapilloma tumour severity), nonlinear covariate functional form (carapace size, sampling year) as well as random effects due to individual heterogeneity and correlation between repeated growth measurements on some turtles. Somatic growth rates were found to be nonlinear functions of carapace size and sampling year but were not a function of low-to-moderate tumour severity. On the other hand, growth rates were significantly lower for turtles with advanced fibropapillomatosis, which suggests a limited or threshold-specific disease effect. However, tumour severity was an increasing function of carapace size—larger turtles tended to have higher tumour severity scores, presumably due to longer exposure of larger (older) turtles to the factors that cause the disease. Hence turtles with advanced fibropapillomatosis tended to be the larger turtles, which confounds size and tumour severity in this study. But somatic growth rates for the Pala’au population have also declined since the mid-1980s (sampling year effect) while disease prevalence and severity increased from the mid-1980s before levelling off by the mid-1990s. It is unlikely that this decline was related to the increasing tumour severity because growth rates have also declined over the last 10–20 years for other green turtle populations resident in Hawaiian waters that have low or no disease prevalence. The declining somatic growth rate trends evident in the Hawaiian stock are more likely a density-dependent effect caused by a dramatic increase in abundance by this once-seriously-depleted stock since the mid-1980s. So despite increasing fibropapillomatosis risk over the last 20 years, only a limited effect on somatic growth dynamics was apparent and the Hawaiian green turtle stock continues to increase in abundance.     

Cambial zones in gills of Bivalvia

Localization and intensity of cell division in the gills of 9 species of Bivalvia were studied. In addition, deoxyribonucleic acid (DNA) synthesis in the gills of two species of mussels (Crenomytilus grayanus and Mytilus edulis) and of the scallop Mizuhopecten yessoensis was investigated by means of radioautography using thymidine-H³. In all bivalves examined, the distribution of branchial cell division was very uneven. Distinctly visible zones exhibited considerably higher mitotic and DNA-synthesizing activities than the remaining branchial epithelium. Topography of these zones appeared similar in all Filibranchia and Eulamellibranchia examined. Areas with enhanced cell division in the gills of Bivalvia are considered as cambial zones or points of growth, apparently serving as a source of growth and/or renewal of the entire organ.     

Oxygen as a limiting factor in phototaxis and in intraclonal spacing of the sea anemone Anthopleura elegantissima

Phototaxis in Anthopleura elegantissima, a sea anemone symbiotic with zooxanthellae, was investigated with special reference to oxygen as a possible controlling factor. Under high oxygen concentrations in seawater, movement towards light was not observed for symbiotic anamones as it was under normal oxygen concentrations. Both aposymbiotic and symbiotic anemones demonstrated movement towards high oxygen concentrations in seawater. Oxygen is, therefore, implicated as a controlling factor in phototaxis. Under laboratory conditions, increased intraclonal spacing occurred with low oxygen concentrations in seawater. In the field, individuals in symbiotic clones were spaced significantly closer than in aposymbiotic clones. Since intraclonal spacing is controlled by oxygen in the laboratory, spacing may also be affected in the field by oxygen; symbiotic clones may be spaced closer because they have better oxygen availability than do aposymbiotic clones.     

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.     

Diel foraging patterns of the sea urchin Centrostephanus coronatus as a predator avoidance strategy

During the day, the diadematid sea urchin Centrostephanus coronatus occupies holes and crevices in shallow subtidal rocky substrata. Individuals emerge from these after sunset and forage on organisms attached to the surrounding rock surface. Each urchin travels <1 m from its shelter and returns to the same one before sunrise. The sheephead wrasse Pimelometopon pulchrum does not remove urchins from their shelters, but will attack and consume urchins placed in normal feeding locations during the daytime. The active periods of the sheephead and the urchin do not overlap; urchins begin foraging about 20 min after the diurnal sheephead retire in the evening and return to their shelters 1 to 2 h before sheephead resume feeding in the morning. We infer that the urchin's daytime crevice-dwelling and nocturnal foraging habits have evolved as a response to sheephead predation. Moreover, because shelters are limited in supply, shelter fidelity may have evolved to insure refuge from sheephead.     

Catch tentacles in the sea anemoneHaliplanella luciae

Evidence is now gathering that indicates that the catch tentacle is an organ of social behavior in sea anemones, which attaches by nematocyst discharge to a feeding tentacle of a neighboring anemone. The typical adhesion response of catch tentacles was observed inHaliplanella luciae and further observations on such behavior were obtained. The active individual that had adhered the tip of an expanded catch tentacle to a neighboring species-mate did not act as a conqueror at all, but rather as a yielder, which would start to run away from the passive neighbor. The final effect on anemone distribution, however, was the same as in cases of aggressive contact in that segregation between individuals was thereby maintained. All individuals tested displayed catch tentacle development as would be predicted by the relative paucity of asexual reproduction in these populations.     

Comparison of larval development and growth of the sea cucumberActinopyga echinites: Ovary-induced ova and DTT-induced ova

AdultActinopyga echinites (Jaeger) were collected from northern Taiwan in September 1989. Oocytes were induced to mature by bathing them in ovary juice (ovary-induced ova) or in 10^–2 M dithiothreitol (DTT-induced ova). The percentage of germinal vesicle breakdown (GVBD) increased from 0.4 to 6.4% in the former treatment and to 84% in the latter. After artificial fertilization, the embryos were cultured in seawater (35 S) at 25 to 28°C. Larvae were fed with the algaIsochysis aff.galbana at 10^4–5 cells/ml. Larvae from ovary-induced ova developed faster (18 d to the doliolaria stage) and grew to a larger size (1.13 mm length) than those from DTT-induced ova (20 d to the doliolaria stage and 0.62 mm in length). On the twelfth day, larvae from ovary-induced ova bear lipid spheres. The number of spheres is positively correlated with larval size. Lipid spheres may provide nutrient reserves for larvae during metamorphosis.     

Effects of food type and ration on reproductive maturation and growth of the sea urchin Strongylocentrotus droebachiensis

We investigated the effects of food quality and quantity on reproductive maturation and growth of juveniles of Strongylocentrotus droebachiensis (Müller) in a 22 month laboratory experiment in which we fed sea urchins four diets: (1) kelp (Laminaria spp.) for 6 d wk⁻¹ and mussel (Mytilus spp.) flesh for 1 d wk⁻¹ (KM); (2) kelp for 7 d wk⁻¹ (high ration, KH); (3) kelp for 1 d wk⁻¹ (low ration, KL) and (4) no food other than encrusting coralline algae (NF). At their first and second opportunity for reproduction, all sea urchins in the KM and KH treatments, and most in the KL treatment were reproductively mature, whereas all sea urchins in the NF treatment remained immature. Gonad index differed significantly among all fed treatments at first and second reproduction, and was highest in the KM and lowest in the KL treatment. Gonad index was similar in both sexes at first reproduction, but it was higher in females than in males at second reproduction. Diet had little or no effect on the relative abundance of spermatocytes, spermatozoa, or nutritive phagocytes in testes at first and second reproduction. In ovaries, nutritive phagocytes were significantly more abundant in females in the KM and KH treatments than in the KL treatment at first reproduction, and significantly more abundant in unfed (NF) than fed (KM, KH, KL) females at second reproduction. Mean oocyte size was similar in all fed females at first reproduction, but significantly larger in fed than unfed females at second reproduction. Mean ovum size was similar in all fed females in both reproductive periods. Increase in test diameter was greatest in the KM treatment and smallest in the KL treatment; sea urchins in the NF treatment decreased slightly in size. Survival was 95 to 100% in all fed sea urchins but significantly lower in unfed ones. The feeding rate on kelp was significantly greater in the KL than the KM and KH treatments. In the KM treatment, the feeding rate on kelp increased significantly over a 6 d period after mussel flesh was provided. Our results demonstrate that a diet of high food quality and quantity accelerates reproductive maturation and growth rate, and enhances gonad production and survival in juvenile and young adult S. droebachiensis. These findings contribute to our understanding of the reproductive ecology of S. droebachiensis in habitats with differing food supplies (e.g., kelp beds and barrens). Our results also can be used to improve aquacultural practices for sea urchins. Received: 3 June 1998 / Accepted: 2 February 1999     

Effects of food ration and feeding regime on growth and reproduction of the sea urchin Strongylocentrotus droebachiensis

To determine the effects food ration and feeding regime on growth and reproduction of Strongylocentrotus droebachiensis (Müller), sea urchins in laboratory aquaria were fed kelp (Laminaria longicruris) supplied at either a high (H, ad libidum daily) or a low (L, ad libidum 1 d wk⁻¹) ration in two successive 12-wk intervals during the reproductive period. After 24 wk, urchins fed the high ration continuously (HH) or for the last 12 wk only (LH) had a significantly greater mean gonad index [(gonad weight/total body weight) × 100] and body weight than urchins fed the low ration continuously (LL) or for the last 12 wk only (HL). Urchins in the HL treatment had a significantly greater gonad index than those in the LL treatment; there was no significant difference in gonad index between the LH and HH treatments. Females had a greater gonad index than males in the low ration (LL and HL) treatments at the end of the experiment; there was no significant difference between sexes in the high ration (LH, HH) treatments. Gametogenesis proceeded to maturation in all treatments and some individuals spawned at the end of the experiment. Females in the high ration (HH and LH) treatments had a greater proportion of nutritive phagocytes in their ovaries than females in the low ration treatments, but there was no effect of feeding treatment on oocyte or ovum size. Feeding treatment had no effect on the relative abundance of nutritive phagocytes in the testes, although the proportion of spermatocytes was higher (and that of spermatozoa lower) in the high ration than in the low ration treatments. Urchins in the high ration treatments had a lower mean jaw height index [(jaw height/test diameter) × 100] and greater mean test diameter than those in the low ration treatments at the end of the experiment, although these differences were not statistically significant. Feeding rate on kelp at the end of the experiment was significantly greater for urchins in the low ration than in the high ration treatments. Our experimental results show that even relatively low rations of kelp support somatic and gonadal growth in S. droebachiensis. Increasing the supply of kelp, particularly during the period of active gametogenesis, results in maximal rates of growth and reproduction. These results suggest that populations of S. droebachiensis in barrens may derive a substantial proportion of their nutrition from drift kelp, which may contribute to their persistence in these habitats. They also explain the large body size, high reproductive effort and fecundity of urchins grazing on kelp beds. These findings have important implications for understanding the dynamics of natural populations of S. droebachiensis and for development of effective aquacultural practices. Received: 17 February 1997 / Accepted: 5 March 1997