Impact of ocean acidification in the metabolism and swimming behavior of the dolphinfish (Coryphaena hippurus) early larvae

Since the industrial revolution, [CO₂]_atm has increased from 280 μatm to levels now exceeding 380 μatm and is expected to rise to 730–1,020 μatm by the end of this century. The consequent changes in the ocean’s chemistry (e.g., lower pH and availability of the carbonate ions) are expected to pose particular problems for marine organisms, especially in the more vulnerable early life stages. The aim of this study was to investigate how the future predictions of ocean acidification may compromise the metabolism and swimming capabilities of the recently hatched larvae of the tropical dolphinfish (Coryphaena hippurus). Here, we show that the future environmental hypercapnia (ΔpH 0.5; 0.16 % CO₂, ~1,600 μatm) significantly (p < 0.05) reduced oxygen consumption rate up to 17 %. Moreover, the swimming duration and orientation frequency also decreased with increasing pCO₂ (50 and 62.5 %, respectively). We argue that these hypercapnia-driven metabolic and locomotory challenges may potentially influence recruitment, dispersal success, and the population dynamics of this circumtropical oceanic top predator.     

Separation of the two Atlantic species of Pomatoceros,P. lamarckii and P. triqueter (Annelida: Serpulidae) by means of biochemical genetics

In order to confirm the status of two proposed species of the serpulid Pomatoceros, the electrophoretic mobilities of 13 enzymes encoded at 19 loci in P. lamarckii (Quatrefages) and P. triqueter (L.) were compared using horizontal starch gel electrophoresis. The serpulids were sampled from Menai Bridge (North Wales), Plymouth (England) and the Isle of Man in the winter of 1981/1982. Six loci were monomorphic or nearly so and showed no useful or significant difference between species. At a further 7 loci, some of the allozymes common to both species were present, though usually at significantly different frquencies, while others were species-specific. The remaining 6 loci presented species-specific allozymes. In P. lamarckii, 61% of the loci were polymorphic compared with 50% in P. triqueter. The mean observed heterozygosity was 0.169 for P. lamarckii and 0.208 for P. triqueter. Calculation of genetic distance and identity gave values of 0.821 and 0.440, respectively. This study, therefore, confirms the existence of two separate species of Pomatoceros as has been previously proposed on the basis of opercular structure and supported by ecological separation where these species occur sympatrically.     

Metal contamination recorded in the sediment of the semi-closed Bakar Bay (Croatia)

This study presents metal levels in the sediments of the Bakar Bay, with its main goal to evaluate recent anthropogenic influence, as well as over previous decades. Sediment profiles at 7 sampling points were taken. Chemical contents in bulk sediment were obtained using ICP, ICP-MS, and AAS methodologies, and 20 most significant elements were presented. Concentrations of selected elements were evaluated by factor statistical analyses to identify their source. Also, metal enrichment factor and geoaccumulation index were calculated, and spatial distribution maps for three sediment layers were constructed. Measured metal concentrations in sediment were compared with concentrations in other sediments from the Adriatic Sea. In addition, a set of sediment quality guidelines were also applied in order to predict the probability of adverse biological effects on the benthic community: This was found not to be very serious. Factor analysis clearly demonstrates the segregation between metals of natural origin resulted from soil and bedrock weathering (Li, Al, Cr, Sc), and with two anthropogenic sources originating from the city of Bakar and bulk cargo terminal (Hg, Pb, Zn, Ag, Sn, and Fe). Mercury (max 0.65 μg g^?1) is found to be the heaviest contaminant, followed by lead (max 71.5 μg g^?1), copper (89.3 μg g^?1), and zinc (156 μg g^?1). However, this study shows that Bakar Bay is considerably less polluted with toxic metals than it was believed.     

Microstructure of the paper nautilus (Argonauta nodosa) shell and the novel application of electron backscatter diffraction (EBSD) to address effects of ocean acidification

Electron backscatter diffraction (EBSD) is a powerful microscopic technique to characterise the crystallography of biomineralisation. Here, we use high-resolution EBSD to characterise one of the least studied shells in the ocean, the female argonaut brood chamber, and to examine the changes in shell microstructure in response to incubation in decreased pH conditions. The thin (225 μm) shell of Argonauta nodosa is magnesium calcite with an average magnesium content of ca. 5.1 Wt % MgCO₃. EBSD and scanning electron microscopy (SEM) revealed that calcification of the shell is bidirectional with formation of irregular crystalline grains. Following a 2 week incubation in a range of pH treatments (pH, 8.1–7.2), shell fragment weight decreased by dissolution in pH ≤ 7.8. EBSD and SEM revealed altered shell crystallography and microstructure at pH ≤ 7.4 due to preferential etching down crystallite grain boundaries and a change in crystalline orientation on both the inner and outer shell surfaces. Our study highlights the value of EBSD for the detailed examination of biogenic carbonates and its potential use in the field of ocean acidification research.     

Populations of the Sydney rock oyster, Saccostrea glomerata, vary in response to ocean acidification

Acidifying oceans are predicted to fundamentally alter marine ecosystems. Over the next century, acute studies suggest that the impacts of climate change on marine organisms and ecosystems may be catastrophic. To date, however, little is known about whether the response of marine organisms varies within a species and whether this provides a potential ??adaptive capacity??. Here, we show that selectively bred lines of the ecologically and economically important estuarine mollusc, the Sydney rock oyster Saccostrea glomerata, are more resilient to ocean acidification than the wild populations. When reared at elevated pCO₂, we found a 25% reduction in shell growth of the selectively bred population of the Sydney rock oyster, Saccostrea glomerata, compared to a 64% reduction in shell growth of wild populations. This study shows that there are significantly different sensitivities to ocean acidification even within the same species, providing preliminary evidence that selective breeding may be a solution for important aquaculture industries to overcome the future effects of ocean acidification.     

Long-term and trans-life-cycle effects of exposure to ocean acidification in the green sea urchin Strongylocentrotus droebachiensis

Anthropogenic CO₂ emissions are acidifying the world’s oceans. A growing body of evidence demonstrates that ocean acidification can impact survival, growth, development and physiology of marine invertebrates. Here, we tested the impact of long-term (up to 16 months) and trans-life-cycle (adult, embryo/larvae and juvenile) exposure to elevated pCO₂ (1,200 μatm, compared to control 400 μatm) on the green sea urchin Strongylocentrotus droebachiensis. Female fecundity was decreased 4.5-fold when acclimated to elevated pCO₂ for 4 months during reproductive conditioning, while no difference was observed in females acclimated for 16 months. Moreover, adult pre-exposure for 4 months to elevated pCO₂ had a direct negative impact on subsequent larval settlement success. Five to nine times fewer offspring reached the juvenile stage in cultures using gametes collected from adults previously acclimated to high pCO₂ for 4 months. However, no difference in larval survival was observed when adults were pre-exposed for 16 months to elevated pCO₂. pCO₂ had no direct negative impact on juvenile survival except when both larvae and juveniles were raised in elevated pCO₂. These negative effects on settlement success and juvenile survival can be attributed to carry-over effects from adults to larvae and from larvae to juveniles. Our results support the contention that adult sea urchins can acclimate to moderately elevated pCO₂ in a matter of a few months and that carry-over effects can exacerbate the negative impact of ocean acidification on larvae and juveniles.     

Antioxidant responses in the polychaete Perinereis gualpensis (Nereididae) exposed to the carbon nanomaterial fullerene (C60)

The objective of this study was to analyse biochemical responses induced by the carbon nanomaterial fullerene (C₆₀) in the polychaete Perinereis gualpensis (Nereididae). The activity of glutathione-S-transferase (GST), glutathione reductase (GR) and glutamate cysteine ligase (GCL), as well as total antioxidant capacity, concentration of glutathione (GSH) and malondialdehyde (TBARS), were analysed. Estuarine worms were maintained in sediments collected at an unpolluted site and spiked with fullerene (3 mg C₆₀·g^?1 sediment). A control group was run in parallel. Scanning electron microscope (SEM) images of sediment and fullerene indicated that the size of the carbon nanomaterial should enable it to be ingested by the polychaete. No evidence of oxidative damage (TBARS) was observed in any of the treatments, and the same was true for GSH and GCL measurements (p>0.05). Total antioxidant capacity was higher in the C₆₀ group after 2 and 7 d when compared with the control group (p<0.05), suggesting that fullerene is acting as an antioxidant. The fact that P. gualpensis is an infaunal organism diminishes the chance of fullerene photoexcitation with consequent reactive oxygen species production. Thus, the data indicated an absence of toxic responses mediated by oxidative stress in estuarine worms exposed to C₆₀ mixed in sediments.     

Predicted impact of ocean acidification on a marine invertebrate: elevated CO<Subscript>2</Subscript> alters response to thermal stress in sea urchin larvae

Ocean acidification (OA) and the biological consequences of altered seawater chemistry have emerged as a significant environmental threat to healthy marine ecosystems. Because a more acidic ocean interferes with fixation of calcium carbonate to form shells or calcified skeletons, future ocean chemistry may significantly alter the physiology of calcifying marine organisms. These alterations may manifest themselves directly in the calcification process, or have synergistic effects with other environmental factors such as elevated temperatures. New tools permit us to explore subtle changes in gene expression patterns in response to environmental conditions. We raised sea urchins (Strongylocentrotus franciscanus) under conditions simulating future atmospheric CO₂ levels of 540 and 970 ppm. When larvae raised under elevated CO₂ conditions were subjected to 1-h acute temperature stress, their ability to mount a physiological response (as measured by expression of the molecular chaperone hsp70) was reduced relative to those raised under ambient CO₂ conditions. These results represent the first use of gene expression assays to study the effects of OA on sea urchin development. They highlight the importance of looking at multiple environmental factors simultaneously as this approach may reveal previously unsuspected biological impacts of atmospheric changes.     

A special adaptation to planktonic life in larvae of the Cassoidea (=Tonnoidea) (Gastropoda)

Living veligers of the Cassoidea have been observed to use a mantle appendage to form and resorb periostracum. Anatomical and histological examinations of a ranellid (Cymatium sp.) larva collected from the Red Sea in 1987 revealed the structure and location of the pallial appendage. The mantle edges of juvenile or adult species of the Cassoidea do not show a comparable specialization. It is demonstrated that cassoid larval conch characters are sufficient to prove the existence of a pallial appendage without anatomical confirmation. A mantle appendage is not known from teleplanic (long-living planktic) veligers of other gastropod superfamilies. In cases where the larval strategies of the latter are known they are totally different. Therefore it is suggested that the adaptation of cassoid larvae to pelagic life is unique among gastropods representing an autapomorphic character of the superfamily.     

Genetic differentiation in two sibling species of the brackish-water polychaete Hediste japonica complex (Nereididae)

Genetic divergence among ten populations of small- and large-egg forms of the brackish-water polychaete Hediste japonica complex was investigated on 14 isozyme loci by electrophoretic analysis. The two forms were distinguishable by complete allele substitutions at five loci, resulting in high genetic differentiation (Nei's D: 0.533 to 0.662). No genetic evidence of hybridization between the two forms was detected in sympatric populations in three rivers. These results indicate that the two forms are reproductively isolated, clearly showing that the two forms are distinct species. The genetic differentiation among populations was higher in the large-egg form (D: 0.005 to 0.111, G _ST: 0.435) than that in the small-egg form (D: 0.000 to 0.001, G _ST: 0.020). This genetic difference between the two forms seems to be attributable to a difference in their life histories. The average expected heterozygosity was low in populations of both the large-egg form (0.005 to 0.068) and the small-egg form (0.014 to 0.038) in comparison with other marine invertebrates. Received: 11 April 1997 / Accepted: 8 September 1997     

A histological study of reproduction in the serpulids<Emphasis Type="Italic"> Pomatoceros triqueter</Emphasis> and<Emphasis Type="Italic"> Pomatoceros lamarckii</Emphasis> (Annelida: Polychaeta)

The reproductive cycles of the serpulid polychaetes Pomatoceros lamarckii (Quatrefages, 1865, as Vermilia) and P. triqueter (Linnaeus, 1767, as Serpula triquetra) were studied by histological examination during 1999 and 2000 at Bantry Bay, south-west Ireland. Gametogenesis, sex ratios and hermaphroditism were also investigated. The male/female ratio in P. lamarckii was approximately equal, but in P. triqueter it deviated significantly in favour of females. There was no significant variation in the sex ratios throughout the year in either species. Results confirm that the species are protandric hermaphrodites. In P. lamarckii, but not in P. triqueter, sexual dimorphism was exhibited in body size, the females being larger than the males of the same species. In both species, the proportion of females increased with increasing body size. Simultaneous hermaphroditism was recorded in P. lamarckii, but not in P. triqueter. There was no clearly defined annual gametogenic cycle in either species. Both species appear to have an extended reproductive season, with numerous small-scale peaks in reproductive maturity that can vary annually. Spawning was broadly synchronous between sexes. Visual observation alone (i.e. without histology) was insufficient to accurately assess reproductive condition in individuals not in, or close to, a ripe state.     

Faecal shield chemical defence is not important in larvae of the tortoise beetle Chelymorpha reimoseri (Chrysomelidae: Cassidinae: Stolaini)

The most frequently investigated defence mechanism among larvae of tortoise beetles is protection by faecal shields, which generally present chemicals that are directly sequestered and/or modified from larval host-plants. In the present work we investigate the tortoise beetle Chelymorpha reimoseri that feeds on the leaves of Ipomoea carnea fistulosa (Convolvulaceae), seeking for the importance of this mechanism to their larvae. We show that 4th instar larvae suffer low predation regardless of the presence of shields in field and laboratory bioassays with ants and chicks. Chloroform extract from larvae without shields provided high protection against predation as shown in bioassays in the field, as well as against chicks, suggesting that C. reimoseri does not rely on the shield for protection. The aqueous extract of the shields did not show activity in such bioassays. The compounds responsible for this protection have yet to be identified, and it remains an open question as to whether they are sequestered from the host plant or de novo biosynthesized. This is the first record of chemical defence in cassidine beetles without the need for faecal shields. These findings indicate that more attention should be paid to chemicals present in the tissues of larvae and/or adults of tortoise beetles; the protective compounds sequestered from host plants or de novo biosynthesized can provide an alternative or complementary strategy against predation in these insects.     

Occurrence and reproductive success of feminized males in the polychaete Capitella capitata (Species Type I)

Natural populations of the cosmopolitan polychaete species, Capitella capitata (Species Type I, Grassle and Grassle 1976) contain males, females and hermaphrodites. Hermaphroditic individuals arise through feminization of males when females are rare. The age-specific survivorships and fecundities of females and hermaphrodites were estimated. There were no significant differences between females and hermaphrodites in survivorship, number of offspring per brood, or percentage of aborted eggs per brood. Net reproductive rates were used to estimate fitness, and the relative fitness of a hermaphrodite as a female ranged from 0.09 to 0.31. The fitness differential was due to the difference in the number of broods that females and hermaphrodites produce. The effects of density, sex ratio, age and body size on the timing of the development of hermaphrodites in groups of siblings were also examined. Hermaphrodites appeared when females were rare or when densities were low. Hermaphrodites never developed in cohorts with larger males unless females were rare. These observations suggest that feminization of males occurs when some males are unable to gain access to females because of mate competition. Feminization does not appear to be correlated with a threshold in body size.     

Metabolic responses of the squat lobster (Pleuroncodes monodon) larvae to low oxygen concentration

Squat lobster populations found in the Humboldt Current System over the continental shelf from ~28 to 37°S release pelagic larvae in sub-surface cold (~11 °C) hypoxic waters. Larvae subsequently spread throughout the water column encountering both normoxic and hypoxic conditions. We analyzed some short- and long-term responses of Pleuroncodes monodon larval metabolism to hypoxia at 11 °C. Routine and postprandial aerobic respiration rates were lower in hypoxia than in normoxia for all zoeal stages. Zoea V oxyconformed, while megalopae oxyregulated down to very low oxygen concentrations. Throughout zoea I development, the rate of nitrogen (protein) accumulation in zoea I was lower, and C:N ratios were higher under hypoxic conditions than in normoxia. Citrate synthase (CS) and malate dehydrogenase (MDH) apparent specific activities (as indicators of aerobic and metabolic potentials, respectively) decreased and remained at the same level, respectively, throughout zoea I reared under hypoxic conditions. Anaerobic to aerobic potential (lactate dehydrogenase (LDH)/CS) was higher in organisms reared under hypoxia, and MDH/LDH potential ratios were characteristic of organisms tolerant to hypoxia. In spite of P. monodon zoea endurance and metabolic adaptations to decreasing oxygen tensions, intense hypoxia as such of their release site would affect their overall condition especially toward the end of the molt cycle. Our results indicate the importance of considering the interaction between environmental oxygen variability and recruitment success.     

Acute and chronic toxicity of tributyltin (TBT) to pediveliger larvae of the bivalve Scrobicularia plana

A static renewal test was run for 30 d to assess the effects of the antifoulant tributyltin (TBT, nominal concentrations of 50, 125, 250 and 500 ng Sn l^-1) on the survival and growth of 240 m-long pediveliger larvae of the bivalve Scrobicularia plana (da Costa) settling in fine sand. Exposure to nominal TBT levels of 125 ng Sn l^-1 (actual range of 37 to 102 ng Sn l^-1, as analysed) resulted in substantial mortalities and negligible shell growth of individuals. More importantly, larval shell growth at the lowest concentration tested (i.e. nominal 50 ng Sn l^-1 or 14 to 32 ng Sn l^-1, as analysed) was significantly reduced (by a factor of 4) and grossly abnormal. In contrast, some control postlarvae had begun developing siphons as they attained a length of 600 to 700 m by the end of this test, thereby completing metamorphosis and becoming juveniles. The results are discussed in relation to the widespread disappearance of S. plana in North Atlantic European estuaries during the 1980s and the concurrent TBT pollution in their waters. It is inferred that TBT is likely to have contributed to the reported demise of clam populations by preventing the successful and timely progress of their pediveliger larvae.     

Settlement behavior and metamorphosis of oyster larvae (Crassostrea gigas) in response to bacterial supernatants

Veliger larvae of the oysterCrassostrea gigas (Thunberg) responded to unknown dissolved chemical inducers found in supernatants of cultures of the bacteriaAlteromonas colwelliana andVibrio cholerae. The response, which was similar to that seen when larvae were exposed to the neurotransmitter precursor L-3,4-dihydroxyphenylalanine (L-DOPA), consisted of an initial settlement phase of swimming with the foot extended and crawling on the substrate. Subsequently larvae attached to the substrate and metamorphosed. The percentage of veligers metamorphosing following inducation of settlement behavior was higher in a group of older larvae, a response similar to that seen with L-DOPA, suggesting that competence to respond to bacterial supernatants is divided into two phases: behavioral competence followed by morphogenetic competence. Following size exclusion chromatography, the molecular weight of the peak containing the activity which induced settlement behavior was determined to be 300 daltons. Autoclaved Marine Broth, which induced low levels of settlement behavior also contained this low molecular weight active peak, suggesting that an oyster settlement inducer is also present in this medium.Contribution # 137 from the Center of Biotechnology, Marine Biotechnology Institute, University of Maryland, USA     

Size regulation in larvae of the crustacean Balanus eburneus (Cirripedia: Thoracica)

We tested the hypothesis that larval size in the acorn barnacle Balanus eburneus Gould (Cirripedia Thoracica) varies in relation to food availability. In March–November 1980, and March–July 1981, larvae were obtained from adult Balanus eburneus collected in the Newport River, North Carolina, USA. Carapace length and width of larvae reared at three different food concentrations were measured. Mean naupliar instar size was independent of food concentration. Mean size of the cypris instar increased with increasing food level. Greater cypris size could be attributed to increased food reserves in the preceding naupliar stage, and was coinciden with inmarked increase in metamorphic success. Variation in instar size remained constant or declined during naupliar development, but increased sharply at the molt to the cyprid. Naupliar size regulation involved: (1) conservation of a molt increment specific for each naupliar-naupliar molt, (2) an inverse relationship between premolt size and the molt increment during the first five naupliar instars, and (3) an increase in the precision of the molt increment at the molt to the sixth naupliar instar. Experimental evidence implies that size regulation in Balanus eburneus limits variation about a fixed final naupliar size (e.g. volume). Measurement of naupliar size, accumulated energy reserves, survival and development time, and cypris metamorphic success indicated that naupliar cuticular growth is the most conservative feature of larval development. The data suggest that maximum naupliar size is limited by escalating metabolic costs during development, while minimum naupliar size is limited by size-related feeding effectiveness.