ALBERT

Description: The clustering of mitochondria near pores in the test walls of foraminifera suggests that these perforations play a critical role in metabolic gas exchange. As such, pore measurements could provide a novel means of tracking changes in metabolic rate in the fossil record. However, in planktonic foraminifera, variation in pore size, density, and porosity have been variously attributed to environmental, biological, and taxonomic drivers, complicating such an interpretation. Here we examine the environmental, biological, and evolutionary determinants of porosity in 718 individuals representing 17 morphospecies of planktonic foraminifera from 6 core tops in the North Atlantic. Using random forest models, we find that porosity is primarily correlated to size and habitat temperature, two key factors in determining metabolic rates. In order to test if this correlation arose spuriously through the association of cryptic species with distinct biomes, we cultured Globigerinoides ruber in three different temperature conditions, and found that porosity increased with temperature. Crucially, these results show that porosity can be plastic: changing in response to environmental drivers within the lifetime of an individual foraminifer. This demonstrates the potential of porosity as a proxy for foraminiferal metabolic rates, with significance for interpreting geochemical data and the physiology of foraminifera in non-analog environments. It also highlights the importance of phenotypic plasticity (i.e., ecophenotypy) in accounting for some aspects of morphological variation in the modern and fossil record.

Description: As anthropogenic activities directly and indirectly increase carbon dioxide (CO2) and decrease oxygen (O2) concentrations in the ocean system, it becomes important to understand how different populations of marine animals will respond. Water that is naturally low in pH, with a high concentration of carbon dioxide (hypercapnia) and a low concentration of oxygen, occurs at shallow depths (200–500 m) in the North Pacific Ocean, whereas similar conditions are absent throughout the upper water column in the North Atlantic. This contrasting hydrography provides a natural experiment to explore whether differences in environment cause populations of cosmopolitan pelagic calcifiers, specifically the aragonitic-shelled pteropods, to have a different physiological response when exposed to hypercapnia and low O2. Using closed-chamber end-point respiration experiments, eight species of pteropods from the two ocean basins were exposed to high CO2 (  800 µatm) while six species were also exposed to moderately low O2 (48 % saturated, or  130 µmol/kg) and a combined treatment of low O2/high CO2. None of the species tested showed a change in metabolic rate in response to high CO2 alone. Of those species tested for an effect of O2, only Limacina retroversa from the Atlantic showed a response to the combined treatment, resulting in a reduction in metabolic rate. Our results suggest that pteropods have mechanisms for coping with short-term CO2 exposure and that there can be interactive effects between stressors on the physiology of these open ocean organisms that correlate with natural exposure to low O2 and high CO2. These are considerations that should be taken into account in projections of organismal sensitivity to future ocean conditions.

Description: Pteropods (pelagic snails) are abundant zooplankton in the Southern Ocean where they are important grazers of phytoplankton, prey for higher trophic levels, and sensitive to environmental change. The Western Antarctic Peninsula (WAP) is a highly dynamic and productive region that has undergone rapid warming, but little is known about how environmental changes there will affect pteropod physiology. In this study, the effects of warming seawater temperatures and shifting food availability on Limacina helicina antarctica metabolism (respiration and excretion) were determined by conducting shipboard experiments that exposed pteropods to a range of temperatures and phytoplankton (food) concentrations. Highest respiration (up to 69 μmol O2/gDW/h) and usually highest excretion rates occurred under higher temperature with more limited metabolic response to food concentration, indicating these factors do not always have an additive effect on pteropod metabolism. The proportion of dissolved organic matter (DOM) to total organic and inorganic dissolved constituents was high and was also significantly affected by shifts in temperature and food. Dissolved organic carbon, nitrogen, and phosphorus (DOC, DON, and DOP) were on average 27, 51, and 11.5% of the total C, N, and P metabolized, respectively. The proportion of total N excreted as DON and the proportion of total P excreted as DOP were significantly affected by a combination of shifting temperature and food concentrations. There were no effects of temperature or food on DOC excretion (mean 8.79 μmol C/gDW/h; range 0.44 to 44) as a proportion of total C metabolized. Metabolic O2:N ratio ranged from 2 to 9 and decreased significantly with increasing temperature and food, indicating a shift toward increased protein catabolism. Metabolic ratios of C, N, and P were all below the canonical Redfield ratio, which has implications for phytoplankton nutrient uptake and bacterial production. Respiration rates at ambient conditions of other WAP pteropods, and excretion rates for Clio pyramidata, were also measured, with respiration rates ranging from 24.39 (Spongiobranchaea australis) to 28.86 (L. h. antarctica) μmol O2/gDW/h. Finally, a CO2 perturbation experiment measuring L. h. antarctica metabolism under pre-industrial and elevated dissolved pCO2 conditions showed no significant change in mean L. h. antarctica respiration or excretion rates with higher pCO2. These insights into the metabolic response of pteropods to ocean variability increase our understanding of the role of zooplankton in biogeochemical cycles and help predict future responses to climate change.