12Thecosome pteropods, a group of calcifying holoplanktonic molluscs, have recently become a 13 research focus due to their potential sensitivity to increased levels of anthropogenic dissolved 14 CO2 in seawater and the accompanying ocean acidification. Some populations, however, already 15 experience high CO2 in their natural distribution during diel vertical migrations. To achieve a 16 better understanding of the mechanisms of pteropod calcification and physiological response to 17 this sort of short duration CO2 exposure, we characterized the gene complement of Clio 18 pyramidata, a cosmopolitan diel migratory thecosome, and investigated its transcriptomic 19 response to experimentally manipulated CO2 conditions. Individuals were sampled from the 20 Northwest Atlantic in the fall of 2011 and exposed to ambient conditions (~380 ppm) and 21 elevated CO2 (~800 ppm, similar to levels experienced during a diel vertical migration) for ~10
Introduction
50The dissolution of anthropogenic carbon dioxide (CO2) into seawater, or ocean acidification 51 (OA), stands to impact a broad variety of marine organisms, especially those that secrete calcium 52 carbonate (CaCO3) shells or skeletons (Fabry et al., 2008). As CO2 equilibrates into seawater, pH 53 and the concentration of carbonate ions decrease, causing CaCO3 to dissociate and affecting the 54 ability of calcifying animals to create and maintain calcareous structures (Gattuso et al., 1999; 55 Riebesell et al., 2000). Furthermore, decreasing pH in the water alters the acid-base balance of 56 the intra-and extracellular fluids of marine organisms (Miles et al., 2007; Seibel and Fabry, 57 2003; Seibel and Walsh, 2001). Since CO2 is produced naturally as a byproduct of respiration, all 58 organisms have physiological mechanisms for maintaining internal cellular pH. These 59 compensatory mechanisms may, however, cease to function if internal acidosis becomes too 60 severe over short time scales and generally fail over extended periods of internal pH imbalance.
61Classical organismal-level metrics (e.g., metabolic rate, calcification, mortality) have revealed a 62 complex pattern of intra-and inter-specific variation in responses to CO2 exposure (reviewed in 63 Hendriks et al., 2010; Kroeker et al., 2013; Kroeker et al., 2010). The variability in these 64 responses, and their dependence on duration of exposure or co-varying stressors, suggest that 65 there is an energetic cost associated with compensating for high CO2 (e.g.,
662011; Langenbuch and Pörtner, 2004; Stumpp et al., 2011b).
68Cummings et al., One group thought to be particularly sensitive to OA is the thecosomatous (i.e., shelled) 71 pteropods. These holoplanktonic gastropods, related to terrestrial snails and also known as sea 72 butterflies, produce thin shells made of aragonite, a highly soluble form of CaCO3. In temperate 73 and polar seas, thecosomes can become a numerically dominant member of the zooplankton 74 community (Hunt et al., 2008). This causes them to be significant consum...