Seawater carbonate chemistry and calcium carbonate of Padina spp., photosynthesis of Padina pavonica in nature CO2 gradients experiment

Predicting the impacts of ocean acidification on coastal ecosystems requires an understanding of the effects on macroalgae and their grazers, as these underpin the ecology of rocky shores. Whilst calcified coralline algae (Rhodophyta) appear to be especially vulnerable to ocean acidification, there is a lack of information concerning calcified brown algae (Phaeophyta), which are not obligate calcifiers but are still important producers of calcium carbonate and organic matter in shallow coastal waters. Here, we compare ecological shifts in subtidal rocky shore systems along CO2 gradients created by volcanic seeps in the Mediterranean and Papua New Guinea, focussing on abundant macroalgae and grazing sea urchins. In both the temperate and tropical systems the abundances of grazing sea urchins declined dramatically along CO2 gradients. Temperate and tropical species of the calcifying macroalgal genus Padina (Dictyoaceae, Phaeophyta) showed reductions in CaCO3 content with CO2 enrichment. In contrast to other studies of calcified macroalgae, however, we observed an increase in the abundance of Padina spp. in acidified conditions. Reduced sea urchin grazing pressure and significant increases in photosynthetic rates may explain the unexpected success of decalcified Padina spp. at elevated levels of CO2. This is the first study to provide a comparison of ecological changes along CO2 gradients between temperate and tropical rocky shores. The similarities we found in the responses of Padina spp. and sea urchin abundance at several vent systems increases confidence in predictions of the ecological impacts of ocean acidification over a large geographical range.

Seawater carbonate chemistry and sea urchin larval size in a laboratory experiment

Ocean acidification results from an increase in the concentrations of atmospheric carbon dioxide (CO2) impacts on marine calcifying species, which is predicted to become more pronounced in the future. By the end of this century, atmospheric pCO2 levels will have doubled relative to the pre-industrial levels of 280 ppm. However, the effects of pre-industrial pCO2 levels on marine organisms remain largely unknown. In this study, we investigated the effects of pre-industrial pCO2 conditions on the size of the pluteus larvae of sea urchins, which are known to be vulnerable to ocean acidification. The larval size of Hemicentrotus pulcherrimus significantly increased when reared at pre-industrial pCO2 level relative to the present one, and the size of Anthocidaris crassispina larvae decreased as the pCO2 levels increased from the pre-industrial level to the near future ones after 3 days' exposure. In this study, it is suggested that echinoid larvae responded to pre-industrial pCO2 levels. Ocean acidification may be affecting some sensitive marine calcifiers even at the present pCO2 level.

Sea urchin response to rising pCO2 shows ocean acidification may fundamentally alter the chemistry of marine skeletons

Ocean acidification caused by an increase in pCO2 is expected to drastically affect marine ecosystem composition, yet there is much uncertainty about the mechanisms through which ecosystems may be affected. Here we studied sea urchins that are common and important grazers in the Mediterranean (Paracentrotus lividus and Arbacia lixula). Our study included a natural CO2 seep plus reference sites in the Aegean Sea off Greece. The distribution of A. lixula was unaffected by the low pH environment, whereas densities of P. lividus were much reduced. There was skeletal degradation in both species living in acidified waters compared to reference sites and remarkable increases in skeletal manganese levels (P. lividus had a 541% increase, A. lixula a 243% increase), presumably due to changes in mineral crystalline structure. Levels of strontium and zinc were also altered. It is not yet known whether such dramatic changes in skeletal chemistry will affect coastal systems but our study reveals a mechanism that may alter inter-species interactions.

Seawater carbonate chemistry, fertilization rate and biological processes of sea urchin Paracentrotus lividus during experiments, 2011

The effect of pH ranging from 8.0 to 6.8 (total scale - pHT) on fertilization, cleavage and larval development until pluteus stage was assessed in an intertidal temperate sea urchin. Gametes were obtained from adults collected in two contrasting tide pools, one showing a significant nocturnal pH decrease (lowest pHT = 7.4) and another where pH was more stable (lowest pHT = 7.8). The highest pHT at which significant effects on fertilization and cleavage were recorded was 7.6. On the contrary, larval development was only affected below pHT 7.4, a value equal or lower than that reported for several subtidal species. This suggests that sea urchins inhabiting stressful intertidal environments produce offspring that may better resist future ocean acidification. Moreover, at pHT 7.4, the fertilization rate of gametes whose progenitors came from the tide pool with higher pH decrease was significantly higher, indicating a possible acclimatization or adaptation of gametes to pH stress.

Bioenergetic trade-offs in the sea cucumber Apostichopus japonicus (Echinodermata: Holothuroidea) in response to CO2-driven ocean acidification

Ocean acidification (OA) caused by excessive CO2 is a potential ecological threat to marine organisms. The impacts of OA on echinoderms are well-documented, but there has been a strong bias towards sea urchins, and limited information is available on sea cucumbers. This work examined the effect of medium-term (60 days) exposure to three pH levels (pH 8.06, 7.72, and 7.41, covering present and future pH variability) on the bioenergetic responses of the sea cucumber, Apostichopus japonicus, an ecologically and economically important holothurian in Asian coasts. Results showed that the measured specific growth rate linearly decreased with decreased pH, leading to a 0.42 %/day decrease at pH 7.41 compared with that at pH 8.06. The impacts of pH on physiological energetics were variable: measured energy consumption and defecation rates linearly decreased with decreased pH, whereas maintenance energy in calculated respiration and excretion were not significantly affected. No shift in energy allocation pattern was observed in A. japonicus upon exposure to pH 7.72 compared with pH 8.06. However, a significant shift in energy budget occurred upon exposure to pH 7.41, leading to decreased energy intake and increased percentage of energy that was lost in feces, thereby resulting in a significantly lowered allocation into somatic growth. These findings indicate that adult A. japonicus is resilient to the OA scenario at the end of the twenty-first century, but further acidification may negatively influence the grazing capability and growth, thereby influencing its ecological functioning as an "ecosystem engineer" and potentially harming its culture output.

Seawater carbonate chemistry and resource allocation and extracellular acid-base status in the sea urchin Strongylocentrotus droebachiensis during experiments, 2012

Anthropogenic CO2 emission will lead to an increase in seawater pCO2 of up to 80-100 Pa (800-1000 µatm) within this century and to an acidification of the oceans. Green sea urchins (Strongylocentrotus droebachiensis) occurring in Kattegat experience seasonal hypercapnic and hypoxic conditions already today. Thus, anthropogenic CO2 emissions will add up to existing values and will lead to even higher pCO2 values >200 Pa (>2000 µatm). To estimate the green sea urchins' potential to acclimate to acidified seawater, we calculated an energy budget and determined the extracellular acid base status of adult S. droebachiensis exposed to moderately (102 to 145 Pa, 1007 to 1431 µatm) and highly (284 to 385 Pa, 2800 to 3800 µatm) elevated seawater pCO2 for 10 and 45 days. A 45 - day exposure to elevated pCO2 resulted in a shift in energy budgets, leading to reduced somatic and reproductive growth. Metabolic rates were not significantly affected, but ammonium excretion increased in response to elevated pCO2. This led to decreased O:N ratios. These findings suggest that protein metabolism is possibly enhanced under elevated pCO2 in order to support ion homeostasis by increasing net acid extrusion. The perivisceral coelomic fluid acid-base status revealed that S. droebachiensis is able to fully (intermediate pCO2) or partially (high pCO2) compensate extracellular pH (pHe) changes by accumulation of bicarbonate (maximum increases 2.5 mM), albeit at a slower rate than typically observed in other taxa (10 day duration for full pHe compensation). At intermediate pCO2, sea urchins were able to maintain fully compensated pHe for 45 days. Sea urchins from the higher pCO2 treatment could be divided into two groups following medium-term acclimation: one group of experimental animals (29%) contained remnants of food in their digestive system and maintained partially compensated pHe (+2.3 mM HCO3), while the other group (71%) exhibited an empty digestive system and a severe metabolic acidosis (-0.5 pH units, -2.4 mM HCO3). There was no difference in mortality between the three pCO2 treatments. The results of this study suggest that S. droebachiensis occurring in the Kattegat might be pre-adapted to hypercapnia due to natural variability in pCO2 in its habitat. We show for the first time that some echinoderm species can actively compensate extracellular pH. Seawater pCO2 values of >200 Pa, which will occur in the Kattegat within this century during seasonal hypoxic events, can possibly only be endured for a short time period of a few weeks. Increases in anthropogenic CO2 emissions and leakages from potential sub-seabed CO2 storage (CCS) sites thus impose a threat to the ecologically and economically important species S. droebachiensis.

Sea-floor videos and photographs (benthos) along 6 shelf profiles from the eastern Weddell Sea, Antarctica taken with remote operated vehicel CHEROKEE during Polarstern cruise ANT-XXI/2

The ROV operations had three objectives: (1) to check, whether the "Cherokee" system is suited for advanced benthological work in the high latitude Antarctic shelf areas; (2) to support the disturbance experiment, providing immediate visual Information; (3) to continue ecological work that started in 1989 at the hilltop situated at the northern margin of the Norsel Bank off the 4-Seasons Inlet (Weddell Sea). The "Cherokee" is was equipped with 3 video cameras, 2 of which support the operation. A high resolution Tritech Typhoon camera is used for scientific observations to be recorded. In addition, the ROV has a manipulator, a still camera, lights and strobe, compass, 2 lasers, a Posidonia transponder and an obstacle avoidance Sonar. The size of the vehicle is 160 X 90 X 90cm. In the present configuration without TMS (tether management system) the deployment has to start with paying out the full cable length, lay it in loops on deck and connect the glass fibres at the tether's spool winch. After a final technical check the vehicle is deployed into the water, actively driven perpendicular to the ship's axis and floatings are fixed to the tether. At a cable length of approx. 50 m, the tether is tightened to the depressor by several cable ties and both components are lowered towards the sea floor, the vehicle by the thruster's propulsion and the depressor by the ship's winch. At 5 m intervals the tether has to be tied to the single conductor cable. In good weather conditions the instruments supporting the navigation of the ROV, especially the Posidonia system, allow an operation mode to follow the ship's course if the ship's speed is slow. Together with the lasers which act as a scale in the images they also allow a reproducible scientific analysis since the transect can be plotted in a GIS system. Consequently, the area observed can be easily calculated. An operation as a predominantly drifting system, especially in areas with bottom near currents, is also possible, however, the connection of the tether at the rear of the vehicle is unsuitable for such conditions. The recovery of the system corresponds to that of the deployment. Most important is to reach the surface of the sea at a safe distance perpendicular to the ship's axis in order not to interfere with the ship's propellers. During this phase the Posidonia transponder system is of high relevance although it has to be switched off at a water depth of approx. 40 m. The minimum personal needed is 4 persons to handle the tether on deck, one person to operate the ship's winch, one pilot and one additional technician for the ROV's operation itself, one scientist, and one person on the ship's bridge in addition to one on deck for whale watching when the Posidonia system is in use. The time for the deployment of the ROV until it reaches the sea floor depends on the water depth and consequently on the length of the cable to be paid out beforehand and to be tightened to the single conductor cable. Deployment and recovery at intermediate water depths can last up to 2 hours each. A reasonable time for benthological observations close to the sea floor is 1 to 3 hours but can be extended if scientifically justified. Preliminary results: after a first test station, the ROV was deployed 3 times for observations related to the disturbance experiment. A first attempt to Cross the hilltop at the northern margin of the Norsel Bank close to the 4- Seasons Inlet was successful only for the first hundreds of metres transect length. The benthic community was dominated in biomass by the demosponge Cinachyra barbata. Due to the strong current of approx. 1 nm/h, the design of the system, and an expected more difficult current regime between grounded icebergs and the top of the hilltop the operation was stopped before the hilltop was reached. In a second attempt the hilltop was successfully crossed because the current and wind situation was much more suitable. In contrast to earlier expeditions with the "sprint" ROV it was the first time that both slopes, the smoother in the northeast and the steeper in the southwest were continuously observed during one cast. A coarse classification of the hilltop fauna shows patches dominated by single taxa: cnidarians, hydrozoans, holothurians, sea urchins and stalked sponges. Approximately 20 % of the north-eastern slope was devastated by grounding icebergs. Here the sediments consisted of large boulders, gravel or blocks of finer sediment looking like an irregularly ploughed field. On the Norsel Bank the Cinachyra concentrations were locally associated with high abundances of sea anemones. Total observation time amounted to 11.5 hours corresponding to almost 6-9 km transect length.

Expression of the Hox gene complex in the indirect development of a sea urchin

Hox complex genes control spatial patterning mechanisms in the development of arthropod and vertebrate body plans. Hox genes are all expressed during embryogenesis in these groups, which are all directly developing organisms in that embryogenesis leads at once to formation of major elements of the respective adult body plans. In the maximally indirect development of a large variety of invertebrates, the process of embryogenesis leads only to a free-living, bilaterally organized feeding larva. Maximal indirect development is exemplified in sea urchins. The 5-fold radially symmetric adult body plan of the sea urchin is generated long after embryogenesis is complete, by a separate process occurring within imaginal tissues set aside in the larva. The single Hox gene complex of Strongylocentrotus purpuratus contains 10 genes, and expression of eight of these genes was measured by quantitative methods during both embryonic and larval developmental stages and also in adult tissues. Only two of these genes are used significantly during the entire process of embryogenesis per se, although all are copiously expressed during the stages when the adult body plan is forming in the imaginal rudiment. They are also all expressed in various combinations in adult tissues. Thus, development of a microscopic, free-living organism of bilaterian grade, the larva, does not appear to require expression of the Hox gene cluster as such, whereas development of the adult body plan does. These observations reflect on mechanisms by which bilaterian metazoans might have arisen in Precambrian evolution.

The Panamic deep sea Echini / by Alexander Agassiz.

v.31 (1904) text

Supplemental report for the sperm cell tests using the sea urchin Arbacia punctulata : training videotape /

Mode of access: Internet.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Sediment rates and structures in Bay of Kiel, Baltic Sea

Density and diversity of bottom fauna population as dependent on sediment types and water depth is largely well known in Kiel Bay. This is in contrast to structures and processes of bioturbation, although generally it has a big influence on the benthic boundary layer and its processes, e.g., the metabolism of the bottom fauna, the mechanical properties, the age dating, and the large field of chemical processes. In the densely inhabited sands and muddy sands of the shallower waters with sediment thicknesses of some decimeters only, bioturbation is usually ubiquitous, and most of the structures left are monotonously of "biodeformational" character. At greater water depths, however, where a sedimentary column of several meters of Holocene is developed, the X-ray radiographs of numerous sediment cores show heterogeneous biogenic structures with regional and stratigraphical differentiation. They are described in terms of ichnofabrics and are interpreted on ethological knowledge of the related macrobenthos species. lmportant organisms creating specific traces include the bivalve Arctica (Cyprina) islandica and the polychaete worm Pectinaria koreni. These species are abundant in Kiel Bay and produce by their crawling-plowing mode of locomotion, a characteristic biogenic stratification, the "plow-sole structure". Other typical biogenic structures are tube traces, which are left by a number of different polychaetes occurring either singly, or as U-pairs mainly in mud sediments. Although sea urchins are rare to absent in Kiel Bay, layers of their characteristic traces Scolicia occur as witness of paleohydrographic events in channel sediments of the central bay. Plow-sole traces, polychaete-tube ichnofabric, Scolicia layers and alternations of laminated and bioturbated layers are considered as building blocks of a future "ichnostratigraphy" of Kiel Bay.

Immunostimulatory effects of different aspects of aquaculture on the host response in the edible sea urchin, paracentrotus lividus

Aquaculture is a fast-growing industry contributing to global food security and sustainable aquaculture, which may reduce pressures on capture fisheries. The overall objective of this thesis was to look at the immunostimulatory effects of different aspects of aquaculture on the host response of the edible sea urchin, Paracentrotus lividus, which are a prized delicacy (roe) in many Asian and Mediterranean countries. In Chapter 1, the importance of understanding the biology, ecology, and physiology of P. lividus, as well as the current status in the culture of this organism for mass production and introducing the thesis objectives for following chapters is discussed. As the research commenced, the difficulties of identifying individuals for repeat sampling became clear; therefore, Chapter 2 was a tagging experiment that indicated PIT tagging was a successful way of identifying individual sea urchins over time with a high tag retention rate. However, it was also found that repeat sampling via syringe to measure host response of an individual caused stress which masked results and thus animals would be sampled and sacrificed going forward. Additionally, from personal observations and discussion with peers, it was suggested to look at the effect that diet has on sea urchin immune function and the parameters I measured which led to Chapter 3. In this chapter, both Laminaria digitata and Mytilus edulis were shown to influence measured immune parameters of differential cell counts, nitric oxide production, and lysozyme activity. Therefore, trials commencing after Trial 5 in Chapter 4, were modified to include starvation in order to remove any effect of diet. Another important aspect of culturing any organism is the study of their immune function and its response to several immunostimulatory agents (Chapter 4). Zymosan A was shown to be an effective immunostimulatory agent in P. lividus. Further work on handled/stored animals (Chapter 5) showed Zymosan A reduced the measured levels of some immune parameters measured relative to the control, which may reduce the amount of stress in the animals. In Chapter 6, animals were infected with Vibrio anguillarum and, although V. anguillarum, impacted immune parameters of P. lividus, it did not cause mortality as predicted. Lastly, throughout this thesis work, it was noted that the immune parameters measured produced different values at different times of the year (Chapter 7); therefore, using collated baseline (control) data, results were compiled to observe seasonal effects. It was determined that both seasonality and sourcing sites influenced immune parameter measurements taken at different times throughout the year. In conclusion, this thesis work fits into the framework of development of aquaculture practices that affect immune function of the host and future research focusing on the edible sea urchin, P. lividus.

Seawater carbonate chemistry and sperm swimming speed, fertilization success of the Australasian sea urchin Heliocidaris erythrogramma in lab experiment

Background: Climate change will lead to intense selection on many organisms, particularly during susceptible early life stages. To date, most studies on the likely biotic effects of climate change have focused on the mean responses of pooled groups of animals. Consequently, the extent to which inter-individual variation mediates different selection responses has not been tested. Investigating this variation is important, since some individuals may be preadapted to future climate scenarios. Methodology/Principal Findings: We examined the effect of CO2-induced pH changes ("ocean acidification") in sperm swimming behaviour on the fertilization success of the Australasian sea urchin Heliocidaris erythrogramma, focusing on the responses of separate individuals and pairs. Acidification significantly decreased the proportion of motile sperm but had no effect on sperm swimming speed. Subsequent fertilization experiments showed strong inter-individual variation in responses to ocean acidification, ranging from a 44% decrease to a 14% increase in fertilization success. This was partly explained by the significant relationship between decreases in percent sperm motility and fertilization success at delta pH = 0.3, but not at delta pH = 0.5. Conclusions and Significance: The effects of ocean acidification on reproductive success varied markedly between individuals. Our results suggest that some individuals will exhibit enhanced fertilization success in acidified oceans, supporting the concept of 'winners' and 'losers' of climate change at an individual level. If these differences are heritable it is likely that ocean acidification will lead to selection against susceptible phenotypes as well as to rapid fixation of alleles that allow reproduction under more acidic conditions. This selection may ameliorate the biotic effects of climate change if taxa have sufficient extant genetic variation upon which selection can act.

Effects of hypercapnia on aspects of feeding, nutrition, and growth in the edible sea urchin Lytechinus variegatus held in culture

Land-based aquaculture facilities experience occasional hypercapnic conditions due to the accumulation of the metabolic waste product carbon dioxide. Pre-gonadal Lytechinus variegatus (horizontal diameter=20 mm) were exposed to control (608 µatm pCO2, pH 8.1) or hypercapnic conditions (1738 µatm pCO2, pH 7.7) in synthetic seawater for 14 weeks. Sea urchins exposed to hypercapnic conditions exhibited significantly slower growth (reduced dry matter production), primarily due to reduced test production. Higher fecal production rates and lower ash absorption efficiency (%) in individuals exposed to hypercapnic conditions suggest the ability to process or retain dietary carbonates may have been affected. Significant increases in neutral lipid storage in the gut and increased soluble protein storage in the gonads of individuals exposed to hypercapnic conditions suggest alterations in nutrient metabolism and storage. Furthermore, organic production and energy allocation increased in the lantern of those individuals exposed to hypercapnic conditions. These results suggest chronic exposure to hypercapnic conditions alters nutrient allocation to organ systems and functions, leading to changes in somatic and reproductive production.