Calcifying echinoid larvae respond to changes in seawater carbonate chemistry with reduced growth and developmental delay. To date, no information exists on how ocean acidification acts on pH homeostasis in echinoderm larvae. Understanding acid-base regulatory capacities is important because intracellular formation and maintenance of the calcium carbonate skeleton is dependent on pH homeostasis. Using H + -selective microelectrodes and the pHsensitive fluorescent dye BCECF, we conducted in vivo measurements of extracellular and intracellular pH (pH e and pH i ) in echinoderm larvae. We exposed pluteus larvae to a range of seawater CO 2 conditions and demonstrated that the extracellular compartment surrounding the calcifying primary mesenchyme cells (PMCs) conforms to the surrounding seawater with respect to pH during exposure to elevated seawater pCO 2 . Using FITC dextran conjugates, we demonstrate that sea urchin larvae have a leaky integument. PMCs and spicules are therefore directly exposed to strong changes in pH e whenever seawater pH changes. However, measurements of pH i demonstrated that PMCs are able to fully compensate an induced intracellular acidosis. This was highly dependent on Na + and HCO 3 − , suggesting a bicarbonate buffer mechanism involving secondary active Na + -dependent membrane transport proteins. We suggest that, under ocean acidification, maintained pH i enables calcification to proceed despite decreased pH e . However, this probably causes enhanced costs. Increased costs for calcification or cellular homeostasis can be one of the main factors leading to modifications in energy partitioning, which then impacts growth and, ultimately, results in increased mortality of echinoid larvae during the pelagic life stage. pH microelectrode | Strongylocentrotus droebachiensis | acid-base regulation | Na + -HCO 3 − transport | epithelial transport S ea urchin larvae have been shown to react with particular sensitivity to CO 2 -induced reductions in seawater pH (1-4). When larvae are chronically exposed to elevated seawater pCO 2 of >0.1 kPa, e.g., as is predicted to occur during the next century in response to anthropogenic CO 2 emissions or through upwelling of low-pH deep water, this sensitivity is reflected in reduced growth and developmental rates (5, 6). Echinoderm larvae are considered to be especially vulnerable to seawater pH reduction and to the associated changes in calcium carbonate saturation state of seawater (Ω Cal ) because their internal skeleton is composed of high magnesium calcite, a highly soluble form of CaCO 3 (7, 8). However, long-term reductions in growth and development might just as well be evoked by other physiological mechanisms that are also sensitive to hypercapnia and the related acid-base disturbances. Recent studies conducted on several marine taxa including mollusks (9) and echinoderms (10) demonstrated increased metabolic rates in response to elevated seawater pCO 2 . It was concluded that reductions in somatic growth and rate of development were caused by a sh...

Anthropogenic CO 2 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 pCO 2 (1,200 latm, compared to control 400 latm) on the green sea urchin Strongylocentrotus droebachiensis. Female fecundity was decreased 4.5-fold when acclimated to elevated pCO 2 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 pCO 2 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 pCO 2 for 4 months. However, no difference in larval survival was observed when adults were pre-exposed for 16 months to elevated pCO 2 . pCO 2 had no direct negative impact on juvenile survival except when both larvae and juveniles were raised in elevated pCO 2 . These negative effects on settlement success and juvenile survival can be attributed to carryover effects from adults to larvae and from larvae to juveniles. Our results support the contention that adult sea urchins can acclimate to moderately elevated pCO 2 in a matter of a few months and that carry-over effects can exacerbate the negative impact of ocean acidification on larvae and juveniles.

Our ability to project the impact of global change on marine ecosystem is limited by our poor understanding on how to predict species sensitivity. For example, the impact of ocean acidification is highly species-specific, even in closely related taxa. The aim of this study was to test the hypothesis that the tolerance range of a given species to decreased pH corresponds to their natural range of exposure. Larvae of the green sea urchin Strongylocentrotus droebachiensis were cultured from fertilization to metamorphic competence (29 days) under a wide range of pH (from pHT  = 8.0/pCO2  ≈ 480 μatm to pHT  = 6.5/pCO2  ≈ 20 000 μatm) covering present (from pHT 8.7 to 7.6), projected near-future variability (from pHT 8.3 to 7.2) and beyond. Decreasing pH impacted all tested parameters (mortality, symmetry, growth, morphometry and respiration). Development of normal, although showing morphological plasticity, swimming larvae was possible as low as pHT  ≥ 7.0. Within that range, decreasing pH increased mortality and asymmetry and decreased body length (BL) growth rate. Larvae raised at lowered pH and with similar BL had shorter arms and a wider body. Relative to a given BL, respiration rates and stomach volume both increased with decreasing pH suggesting changes in energy budget. At the lowest pHs (pHT  ≤ 6.5), all the tested parameters were strongly negatively affected and no larva survived past 13 days post fertilization. In conclusion, sea urchin larvae appeared to be highly plastic when exposed to decreased pH until a physiological tipping point at pHT  = 7.0. However, this plasticity was associated with direct (increased mortality) and indirect (decreased growth) consequences for fitness.

No abstract

Abstract:This study investigated the effects of seawater pH (i.e. 8.10, 7.85 and 7.60) and temperature (16 and 19°C) on (i) the abiotic conditions in the fluid surrounding the embryo (viz. the perivitelline fluid),(ii) growth, development and (iii) cuttlebone calcification of embryonic and juvenile stages of the cephalopod Sepia officinalis. Egg swelling increased in response to acidification or warming, leading to an increase in egg surface while the interactive effects suggested a limited plasticity of the swelling modulation. Embryos experienced elevated pCO 2 conditions in the perivitelline fluid (> 3-fold higher pCO 2 than that of ambient seawater), rendering the medium under-saturated even under ambient conditions. The growth of both embryos and juveniles was unaffected by pH, whereas 45 Ca incorporation in cuttlebone increased significantly with decreasing pH at both temperatures. This phenomenon of hypercalcification is limited to only a number of animals but does not guarantee functional performance and calls for better mechanistic understanding of calcification processes.

Diurnal variability of ocean CO2 system variables is poorly constrained. Here, this variability and its drivers are assessed using 3‐h observations collected over 8–140 months at 37 stations located in diverse marine environments. Extreme diurnal variability, that is, when the daily amplitude exceeds the 99th percentile of diurnal variability, is comparable in magnitude to the seasonal amplitude and can surpass projected changes in mean states of pCO2 and [H+] over the twenty‐first century. At coastal sites and near coral reefs, extremes in diurnal amplitudes reach 187 ± 85 and 149 ± 106 μatm for pCO2, 0.21 ± 0.08 and 0.11 ± 0.07 for pH, and 1.2 ± 0.5 and 0.8 ± 0.4 for Ωarag, respectively. Extreme diurnal variability is weaker in the open ocean, but still reaches 47 ± 18 μatm for pCO2, 0.04 ± 0.01 for pH, and 0.25 ± 0.11 for Ωarag. Diurnal variability of the ocean CO2 system is considerable and likely to respond to increasing CO2. Therefore, it should be represented in Earth system models.

Along the Norwegian coasts and margins, extensive reefs of the stony coral Lophelia pertusa act as hotspots for local biodiversity. Climate models project that the temperature of Atlantic deep waters could rise by 1-3 • C by 2100. In this context, understanding the effects of temperature on the physiology of cold-water species will help in evaluating their resilience to future oceanic changes. We investigated the response of L. pertusa to stepwise short-term increases in temperature. We sampled corals from four reefs, two located north of the Arctic circle and two at the mid-Norwegian shelf (boreal). In on-board experiments (one per reef), the sampled fragments were exposed to increasing temperatures from 5 to 15 • C over 58 h. Respiration increased linearly by threefold for a 10 • C increase. The short-term temperature increase did not induce mortality, cellular (neutral red assay for lysosome membrane stability; but one exception) or oxidative stress (lipid peroxidation assay) -to a few exceptions. However, the variability of the respiration responses depended on the experiment (i.e., reef location), possibly linked to the genetic structure of the individuals that we sampled (e.g., clones or siblings). The corals from the Arctic and boreal regions appear to have a high tolerance to the rapid temperature fluctuations they experience in the field. Over extended periods of time however, an increased metabolism could deplete the energy stored by the corals, if not met by an increased food availability and/or uptake. Empirical data on organisms' thermal performance curves, such as the one presented in this study for L. pertusa, will be useful to implement predictive models on the responses of species and populations to climate change.

Ocean acidification (OA) is increasing due to anthropogenic CO2 emissions and poses a threat to marine species and communities worldwide. To better project the effects of acidification on organisms’ health and persistence, an understanding is needed of the 1) mechanisms underlying developmental and physiological tolerance and 2) potential populations have for rapid evolutionary adaptation. This is especially challenging in nonmodel species where targeted assays of metabolism and stress physiology may not be available or economical for large-scale assessments of genetic constraints. We used mRNA sequencing and a quantitative genetics breeding design to study mechanisms underlying genetic variability and tolerance to decreased seawater pH (-0.4 pH units) in larvae of the sea urchin Strongylocentrotus droebachiensis. We used a gene ontology-based approach to integrate expression profiles into indirect measures of cellular and biochemical traits underlying variation in larval performance (i.e., growth rates). Molecular responses to OA were complex, involving changes to several functions such as growth rates, cell division, metabolism, and immune activities. Surprisingly, the magnitude of pH effects on molecular traits tended to be small relative to variation attributable to segregating functional genetic variation in this species. We discuss how the application of transcriptomics and quantitative genetics approaches across diverse species can enrich our understanding of the biological impacts of climate change.