Comments on and implications of a steady-state in coastal marine ecosystems

Coastal ecosystems can be thought of as being established by a number of physico-geochemical drivers, e.g. geochemistry and bathymetry of the basins, climate, tidal and freshwater flows, natural and anthropogenic inputs of nutrients and toxins, all of which exert an influence on the resulting communities of organisms. Depending on the interactions among the major drivers, ecosystems may occur on both large and small scales and be basin-wide or within basins. For individual and separate ecosystems to exist with some permanence in time, e.g. reach a steady-state, they also have to be ‘defended’. Defences are mechanisms that counter changes to maintain the status quo. We argue, and present evidence to support the notion, that the defence mechanisms are inextricably tied to primary production and the biogeochemical cycling of organic matter and provide buffers that mitigate potentially adverse impacts by trace toxins. Colloid pumping, production of complexing ligands and sulfide formation are some of the mechanisms that control trace substances. Current methods for assessing ecosystems do not address the issue of steady-state, nor do they take account of defence activities, e.g. buffering. Therefore, they cannot assess the ‘robustness’ of ecosystems or their ability to resist change, for good or bad. Also, defence mechanisms may, for a time, mask future potentially serious impacts, suggesting that monitoring efforts with limited budgets should consider the measurement of the inputs into ecosystems as well as the immediate or short-term result of the inputs.     

Biological benthic tools as indicators of coastal marine ecosystems health

Coastal marine ecosystems are increasingly subjected to environmental stress and degradation due to pollution. Several research programmes have addressed this problem and produced relevant data sets for specific areas, often including consistent sets of environmental and biological variables. The value of existing information gathered from these types of data can be largely increased by combining them into a common data set to determine globally applicable relationships. To perform this exercise, the Intergovernmental Oceanographic Commission (IOC) of UNESCO has recently formed the Ad hoc Study Group on Benthic Indicators (http://www.ioc.unesco.org/benthicindicators) with the aim of developing robust indicators of benthic health. In this paper, initial products and ongoing activities of this international initiative are described and discussed. An expansion of initial IOC/UNESCO research on benthic fauna-organic carbon relationships is also presented. As part of this follow-up research, the relationship between total organic carbon concentrations of sediment and abundance, biomass and species diversity of benthic macrofauna was evaluated using data sets from 2 different regions of the world comprising 3 different coastal marine environments. The ability of identifying threshold levels in selected variables that could serve as indicators of related adverse environmental conditions leading to stress in the benthos is envisaged within the frame of a larger joint analysis, carried out by the IOC/UNESCO Study Group on Benthic Indicators, of merged data sets from several coastal regions worldwide.     

Biological benthic tools as indicators of coastal marine ecosystems health

Coastal marine ecosystems are increasingly subjected to environmental stress and degradation due to pollution. Several research programmes have addressed this problem and produced relevant data sets for specific areas, often including consistent sets of environmental and biological variables. The value of existing information gathered from these types of data can be largely increased by combining them into a common data set to determine globally applicable relationships. To perform this exercise, the Intergovernmental Oceanographic Commission (IOC) of UNESCO has recently formed the Ad hoc Study Group on Benthic Indicators (http://www.ioc.unesco.org/benthicindicators) with the aim of developing robust indicators of benthic health. In this paper, initial products and ongoing activities of this international initiative are described and discussed. An expansion of initial IOC/UNESCO research on benthic fauna-organic carbon relationships is also presented. As part of this follow-up research, the relationship between total organic carbon concentrations of sediment and abundance, biomass and species diversity of benthic macrofauna was evaluated using data sets from 2 different regions of the world comprising 3 different coastal marine environments. The ability of identifying threshold levels in selected variables that could serve as indicators of related adverse environmental conditions leading to stress in the benthos is envisaged within the frame of a larger joint analysis, carried out by the IOC/UNESCO Study Group on Benthic Indicators, of merged data sets from several coastal regions worldwide.     

Oceanic variability and coastal topography shape genetic structure in a long-dispersing sea urchin

Understanding the scale of marine population connectivity is critical for the conservation and sustainable management of marine resources. For many marine species adults are benthic and relatively immobile, so patterns of larval dispersal and recruitment provide the key to understanding marine population connectivity. Contrary to previous expectations, recent studies have often detected unexpectedly low dispersal and fine-scale population structure in the sea, leading to a paradigm shift in how marine systems are viewed. Nonetheless, the link between fine-scale marine population structure and the underlying physical and biological processes has not been made. Here we show that patterns of genetic structure and population connectivity in the broadcast-spawning and long-distance dispersing sea urchin Centrostephanus rodgersii are influenced by physical oceanographic and geographic variables. Despite weak genetic differentiation and no isolation-by-distance over thousands of kilometers among samples from eastern Australia and northern New Zealand, fine-scale genetic structure was associated with sea surface temperature (SST) variability and geography along the southeastern Australian coast. The zone of high SST variability is characterized by periodic shedding of eddies from the East Australian Current, and we suggest that ocean current circulation may, through its influence on larval transport and recruitment, interact with the genetic consequences of large variance in individual reproductive success to generate patterns of fine-scale patchy genetic structure. If proven consistent across species, our findings suggest that the optimal scale for fisheries management and reserve design should vary among localities in relation to regional oceanographic variability and coastal geography.     

Effects of ethynylestradiol and bisphenol A on the development of sea urchin embryos and juveniles

The effects of various concentrations of a synthetic estrogen, ethynylestradiol (EER), and an estrogenic compound, bisphenol A (BPA), on the development of two sea urchin species, Hemicentrotus pulcherrimus and Strongylocentrotus nudus, were examined. At concentrations of 2.5 μM or higher of EER the zygotes did not hatch; at 0.98–1.25 μM they developed but had abnormal morphology; and at concentrations lower than 1 μM there was no effect on embryogenesis. The dose dependency was the same for both species, but more embryos of S. nudus showed exogastrulation at 3–10 μM EER. The effect of BPA on early development was less remarkable than that of EER; most embryos developed normally even in the presence of 2.5 μM BPA. The feeding larvae progressed and finally metamorphosed into juveniles even at 0.1–3 μM EER. The chemicals had opposing effects on the growth of the juveniles. A low dose of EER (0.1–1 μM) promoted growth and the average diameter of the test was bigger than that of the control group; the same dose of BPA suppressed growth and the test was smaller than that of the control group. These results show that the sensitivity and response to endocrine disrupter chemicals changes markedly during the ontogeny of sea urchins. Physical and Chemical Impacts on Marine Organisms, a Bilateral Seminar Italy-Japan held in November 2004     

Effects of ultraviolet-B enhancement on marine trophic levels in a stratified coastal system

We examined the effects of enhanced UV-B radiation (relative to ambient) on marine trophic levels inhabiting a stratified coastal ecosystem, using living models (13 000 liter marine enclosures) of a temperate estuarine water column. The experiment was carried out in June and July 1994 on a plankton community drawn from lower Narragansett Bay, Rhode Island, USA. The effects of altered UV-B radiation (elevated 50% over ambient, tenfold DNA-weighted) on three trophic strata: the primary producers (photosynthetic algae), primary herbivores (copepods), and fish eggs and larvae (Anchoa mitchilli Cuvier and Valenciennes) were examined. The goal was to determine if UV-B–induced alterations at the base of the food chain had impacts on other elements of the trophic web. Phytoplankton abundance (P=0.02) and biomass (P=0.007) were significantly reduced in UV-B–enhanced treatments, above but not below the thermocline (2.25 m), during the second week of the study. Copepod nauplii were significantly less abundant in UV-B–enhanced mesocosms than in control treatments during the third and fourth weeks of the study (P=0.01). A portion of the impact on nauplii may be a result of alterations at the base of the food web. The greater mortality of nauplii in UV-B–enhanced systems did not translate to reduced abundance of copepodite (P=0.83) or adult (P=0.29) copepods. No significant effects were observed for microzooplankton (P=0.15). Neither the mortality rates nor the growth rates of larval anchovy were significantly affected by the experimental increase in UV-B (P>0.05). Despite the tenfold increase in biologically damaging UV-B, effects were not seen at higher trophic levels, most likely because of the rapid extinction of UV-B in the highly colored coastal water. Received: 13 November 1996 / Accepted: 18 July 1997     

Effects of predators on sea urchin density and habitat use in a southern California kelp forest

Grazing sea urchins can reduce kelp abundance and therefore strongly affect kelp forest community structure. Despite the ecological importance of sea urchins, direct field studies on the role that urchin predators play in shaping urchin populations are rare for southern California. We conducted surveys and manipulative experiments within kelp forests near San Diego, CA, (32–51′28″N, 117–16′00″W) from 2006 to 2009 to determine whether predators such as sheephead (Semicossyphus pulcher) and spiny lobsters (Panulirus interruptus) may be linked to purple urchin (Strongylocentrotus purpuratus) and red urchin (Strongylocentrotus franciscanus) distribution and habitat use, as well as purple urchin density-dependent mortality. Purple urchins were less dense and more cryptic inside a local marine protected area (MPA) that contained high predator abundance than in nearby heavily fished areas, whereas red urchins rarely were found outside the MPA. Urchin proportional mortality was inversely density dependent during the day when sheephead were active, despite fish aggregations in plots of high urchin density, but was density independent during the night when lobsters were active. Urchin mortality was reduced under understory algal cover during the day, but not during the night. Examining whether urchin mortality from predation is density dependent and how habitat complexity influences this relationship is imperative because behavioral changes and increases in urchin populations can have vast ecological and economic consequences in kelp forest communities.     

Limited effects of Antarctic ozone depletion on sea urchin development

The sea urchin, Sterechinus neumayeri, has a circumpolar distribution and is an abundant species in benthic communities of the Antarctic. Reproduction occurs during austral spring, when ozone concentrations over the past 25 years have been reduced by 50% or more, potentially exposing the planktonic embryos and larvae to elevated levels of UVB. During spring of 1996, cultures of S. neumayeri embryos incubated under ambient and partitioned sunlight (minus UVB) at static depths between 0 and 7 m were analyzed for DNA damage [cyclobutane pyrimidine dimers (CPDs)] and morphological abnormalities. At 0-m and 1-m depths, nearly 100% of embryos developed abnormally, even under UVB-shielded conditions where little or no DNA damage accumulated. At depths >3 m, reduced or no abnormality was evident and DNA damage was negligible. Although UVB contributed to 0–65% of solar-induced abnormalities, the mean contribution was 11±17% and UVB was not primarily responsible for observed defects in urchin development. Moreover, developmental responses were not linearly related to ambient UVB gradients as might be expected, but are better characterized relative to threshold levels of total UVB exposure. Accumulated exposures of 25 kJ m^–2 ambient UVB caused minimal DNA damage and allowed normal embryological development to proceed. Higher UVB exposures (especially 80 kJ m^–2) precluded normal development. An ancillary threshold limit of 17 CPDs mb^–1 has been identified as the level of DNA damage that proscribes abnormal development. While higher wavelengths of UVA and visible light are not affected by ozone concentration and do not initiate significant CPD DNA damage, they did interfere significantly with the embryological development of S. neumayeri. It is concluded that exposure to increased UVB during recent Antarctic ozone-depletion cycles probably has only a small degree of impact relative to the magnitude of other solar effects on the developmental success of Sterechinus embryos, or compared to spawning seasons before ozone depletion (i.e., years prior to 1978).Communicated by P.W. Sammarco, Chauvin     

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     

Development of the echinothurid sea urchin Asthenosoma ijimai

Development of an echinothurid sea urchin is described for the first time; eggs of Asthenosoma ijimai (Yoshiwara) have been fertilized in the laboratory, and development has been followed light-microscopically up to the early juvenile stage. The eggs, which are orange and float, are 1.2 mm in diameter, by far the largest echinoid eggs on record. The embryos, larvae and early juveniles are lecithotrophic, and no exogenous food is needed for development. The embryonic stages include a remarkable biscuit-shaped late blastula, which has never been described for any echinoid before. The larva, with its para-arms unsupported by skeletal ossicles, is unlike the echinoid prism or pluteus and more resembles the early bipinnaria or early auricularia of asteroids and holothurians, respectively. All stages through late larvae float just beneath the surface until settlement occurs during the third and fourth weeks at 20°C. Metamorphosis from late larva to juvenile is gradual and no part of the larval body appears to be cast off.This paper is dedicated to Professor Juro Ishida on the occasion of his seventieth birthday.     

Antigenic specificity of the sea urchin sperm surface

The antigenic composition of the sperm surface in Paracentrotus lividus Lam. and Sphaerechinus granularis Lam. was studied using the sperm agglutination test with homologous and heterologous antisera. The results indicated that sperm of both species have at least one antigenic determinant located on their surface and shared by both species. In addition, Paracentrotus sperm bear at least one species specific determinant on their surface. The existence of antigenic determinants common to gametes of both species was demonstrated.     

Self-purification capacity and management of Baltic coastal ecosystems

It is expected that the Baltic region becomes a major centre of economic growth and prosperity in Europe already during this decade (Anon. 2000). Therefore, an Agenda 21 for the Baltic region (Baltic 21) was developed to ensure a sustainable development. Especially the coastal ecosystems are subject to increasing anthropogenic pressure e.g. eutrophication, traffic, harbours, tourism or offshore wind parks. Eutrophication remains the main ecological problem in the Baltic Sea and has serious negative social and economical consequences. Inner and outer coastal waters play an important role as buffers and filters for the Baltic proper. Consequently, the utilization and preservation of their self-purification capacity is of great importance. Combined results of our own coastal research and of the international workshop ‘Baltic coastal ecosystems: structure, function and coastal zone management’ (Rostock University, November 2001) are presented here. Conclusions for an improved integrated coastal zone management of Baltic coastal ecosystems will be presented.     

Impacts of UV-B radiation on short-term cultures of sea urchin coelomocytes

Three specialized cell types constitute the heterogeneous population present in the coelomic cavity fluid of sea urchins. The list includes: phagocytes, which undergo a stress-induced petaloid-filopodial transition, white or red amoebocytes and vibratile cells. As a whole, they act as the immune defense system of the sea urchin and respond to environmental and experimental challenge triggering specific stress markers. Here we extended our studies on coelomocytes short-term cultures by describing the morphology and occurrence of each cell type and analyzing their response to UV-B radiation at the biochemical level and with respect to DNA damage. The effects of different doses, ranging from 500 to 2,000 J/m², on cells, which were then cultured in vitro for 1–6 h were tested. As early as 1 h after irradiation we found an increase in the levels of the heat shock protein70 (hsp) stress marker. A peak at 1,000 J/m² corresponding to a 3.4-fold hsp70 increase over the levels of control coelomocytes was observed. Concurrently, we found an increase in the number of apoptotic nuclei detected by Hoechst staining, which varied from 7.1 to 15.6% in cells that were exposed to 500 and 2,000 J/m², respectively. On the basis of our findings we confirm sea urchin coelomocytes as a sensitive cell culture system for environmental studies and propose their use for the analysis on the effects of UV-B radiation. Physical and Chemical Impacts on Marine Organisms, a Bilateral Seminar Italy-Japan held in November 2004. This paper is dedicated to the memory of Aurelia Koros, Ph. D., University of Pittsburg, who committed her scientific life to the study of natural killer cell antigens expressed in small lung carcinoma cells and sea urchin coelomocytes.     

Killing of marine phytoplankton by a gliding bacterium Cytophaga sp., isolated from the coastal sea of Japan

A marine gliding bacterium Cytophaga sp. (strain J18/M01) was isolated from Harima-Nada, eastern Seto Inland Sea, Japan in 1990. This bacterium preys upon various species of marine phytoplankton. All of the five raphidophycean flagellates, all of the four diatoms, and one of the two dinoflagellates examined were killed within a few days when cultured with the bacterium. The bacterium presumably achieves this by direct attack, because the culture filtrate in which host organisms were totally destroyed had no significant effects on the growth of the same host organism (Chattonella antiqua). If one or a few bacterial cells were inoculated into C. antiqua culture, all of the host organisms were killed. The bacterium proliferated in filter-sterilized seawater, suggesting its ubiquitous existence in the coastal sea. The killing of phytoplankton by bacteria such as Cytophaga sp. J18/M01 may be a significant factor influencing the population dynamics of phytoplankton in nature and may contribute to the sudden disappearance of red tides in the coastal sea. Bacterial destruction of phytoplankton may also be a factor that regulates primary productivity in marine ecosystems.     

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     

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.     

Culture experiments on the sea urchin Strongylocentrotus pulcherrimus fed an artificial diet

The omnivorous sea urchin Strongylocentrotus pulcherrimus inhabits the coast of Japan. It also ingests foreign foods, which are not normally present in the natural habitat. In cultures, foreign foods, such as white fish meal, shellfish meal, milk casein, gelatin, soybean meal, yellow corn, alfalfa and yeast were provided in the form of a moist and soft material and their ingestion and digestibility were examined. The foreign foods were prepared by mixing with agar-agar. Digestibility of the diet, and that of nitrogen contained in the diet, was estimated as 54.7 and 64.7% dry weight. Sea urchins cultivated over a period of 5 months on such a foreign diet exhibited 24% increase in wet weight during this period. Occasionally, the ingestion of artificial food suddenly decreased, but was soon resumed upon supplying a hot-water extract of brown algae.