Abstract. Future ocean acidification has the potential to adversely affect many marine organisms. A growing body of evidence suggests that many species could suffer from reduced fertilization success, decreases in larval-and adult growth rates, reduced calcification rates, and even mortality when being exposed to near-future levels (year 2100 scenarios) of ocean acidification. Little research focus is currently placed on those organisms/taxa that might be less vulnerable to the anticipated changes in ocean chemistry; this is unfortunate, as the comparison of more vulnerable to more tolerant physiotypes could provide us with those physiological traits that are crucial for ecological success in a future ocean. Here, we attempt to summarize some ontogenetic and lifestyle traits that lead to an increased tolerance towards high environmental pCO 2 . In general, marine ectothermic metazoans with an extensive extracellular fluid volume may be less vulnerable to future acidification as their cells are already exposed to much higher pCO 2 values (0.1 to 0.4 kPa, ca. 1000 to 3900 µatm) than those of unicellular organisms and gametes, for which the ocean (0.04 kPa, ca. 400 µatm) is the extracellular space. A doubling in environmental pCO 2 therefore only represents a 10% change in extracellular pCO 2 in some marine teleosts. High extracellular pCO 2 values are to some degree related to high metabolic rates, as diffusion gradients need to be high in order to excrete an amount of CO 2 that is directly proportional to the amount of O 2 consumed. In active metazoans, such as teleost fish, cephalopods and Correspondence to: F. Melzner (fmelzner@ifm-geomar.de) many brachyuran crustaceans, exercise induced increases in metabolic rate require an efficient ion-regulatory machinery for CO 2 excretion and acid-base regulation, especially when anaerobic metabolism is involved and metabolic protons leak into the extracellular space. These ion-transport systems, which are located in highly developed gill epithelia, form the basis for efficient compensation of pH disturbances during exposure to elevated environmental pCO 2 . Compensation of extracellular acid-base status in turn may be important in avoiding metabolic depression. So far, maintained "performance" at higher seawater pCO 2 (>0.3 to 0.6 kPa) has only been observed in adults/juveniles of active, high metabolic species with a powerful ion regulatory apparatus. However, while some of these taxa are adapted to cope with elevated pCO 2 during their regular embryonic development, gametes, zygotes and early embryonic stages, which lack specialized ion-regulatory epithelia, may be the true bottleneck for ecological success -even of the more tolerant taxa.Our current understanding of which marine animal taxa will be affected adversely in their physiological and ecological fitness by projected scenarios of anthropogenic ocean acidification is quite incomplete. While a growing amount of empirical evidence from CO 2 perturbation experiments suggests that several taxa might react quite se...

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BackgroundBiogeochemical elemental cycling is driven by primary production of biomass via phototrophic phytoplankton growth, with 40% of marine productivity being assigned to diatoms. Phytoplankton growth is widely limited by the availability of iron, an essential component of the photosynthetic apparatus. The oceanic diatom Thalassiosira oceanica shows a remarkable tolerance to low-iron conditions and was chosen as a model for deciphering the cellular response upon shortage of this essential micronutrient.ResultsThe combined efforts in genomics, transcriptomics and proteomics reveal an unexpected metabolic flexibility in response to iron availability for T. oceanica CCMP1005. The complex response comprises cellular retrenchment as well as remodeling of bioenergetic pathways, where the abundance of iron-rich photosynthetic proteins is lowered, whereas iron-rich mitochondrial proteins are preserved. As a consequence of iron deprivation, the photosynthetic machinery undergoes a remodeling to adjust the light energy utilization with the overall decrease in photosynthetic electron transfer complexes.ConclusionsBeneficial adaptations to low-iron environments include strategies to lower the cellular iron requirements and to enhance iron uptake. A novel contribution enhancing iron economy of phototrophic growth is observed with the iron-regulated substitution of three metal-containing fructose-bisphosphate aldolases involved in metabolic conversion of carbohydrates for enzymes that do not contain metals. Further, our data identify candidate components of a high-affinity iron-uptake system, with several of the involved genes and domains originating from duplication events. A high genomic plasticity, as seen from the fraction of genes acquired through horizontal gene transfer, provides the platform for these complex adaptations to a low-iron world.

Progressive ocean acidification due to anthropogenic CO2 emissions will alter marine ecosytem processes. Calcifying organisms might be particularly vulnerable to these alterations in the speciation of the marine carbonate system. While previous research efforts have mainly focused on external dissolution of shells in seawater under saturated with respect to calcium carbonate, the internal shell interface might be more vulnerable to acidification. In the case of the blue mussel Mytilus edulis, high body fluid pCO2 causes low pH and low carbonate concentrations in the extrapallial fluid, which is in direct contact with the inner shell surface. In order to test whether elevated seawater pCO2 impacts calcification and inner shell surface integrity we exposed Baltic M. edulis to four different seawater pCO2 (39, 142, 240, 405 Pa) and two food algae (310–350 cells mL−1 vs. 1600–2000 cells mL−1) concentrations for a period of seven weeks during winter (5°C). We found that low food algae concentrations and high pCO2 values each significantly decreased shell length growth. Internal shell surface corrosion of nacreous ( = aragonite) layers was documented via stereomicroscopy and SEM at the two highest pCO2 treatments in the high food group, while it was found in all treatments in the low food group. Both factors, food and pCO2, significantly influenced the magnitude of inner shell surface dissolution. Our findings illustrate for the first time that integrity of inner shell surfaces is tightly coupled to the animals' energy budget under conditions of CO2 stress. It is likely that under food limited conditions, energy is allocated to more vital processes (e.g. somatic mass maintenance) instead of shell conservation. It is evident from our results that mussels exert significant biological control over the structural integrity of their inner shell surfaces.

Abstract. CO 2 emissions are leading to an acidification of the oceans. Predicting marine community vulnerability towards acidification is difficult, as adaptation processes cannot be accounted for in most experimental studies. Naturally CO 2 enriched sites thus can serve as valuable proxies for future changes in community structure. Here we describe a natural analogue site in the Western Baltic Sea. Seawater pCO 2 in Kiel Fjord is elevated for large parts of the year due to upwelling of CO 2 rich waters. Peak pCO 2 values of >230 Pa (>2300 µatm) and pH NBS values of <7.5 are encountered during summer and autumn, average pCO 2 values are ∼70 Pa (∼700 µatm). In contrast to previously described naturally CO 2 enriched sites that have suggested a progressive displacement of calcifying auto-and heterotrophic species, the macrobenthic community in Kiel Fjord is dominated by calcifying invertebrates. We show that blue mussels from Kiel Fjord can maintain control rates of somatic and shell growth at a pCO 2 of 142 Pa (1400 µatm, pH NBS = 7.7). Juvenile mussel recruitment peaks during the summer months, when high water pCO 2 values of ∼100 Pa (∼1000 µatm) prevail. Our findings indicate that calcifying keystone species may be able to cope with surface ocean pH NBS values projected for the end of this century when food supply is sufficient. However, owing to non-linear synergistic effects of future acidification and upwelling of corrosive water, peak seawater pCO 2 in Kiel Fjord and many other productive estuarine habitats could increase to values >400 Pa (>4000 µatm). These changes will most likely affect calcification and recruitment, and increase external shell dissolution.

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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...

Calcification in coccolithophores has high energy demand but brings multiple benefits enabling diversity of ecology and form.