Calcifying phytoplankton in natural laboratories for understanding ocean acidification

Abstract

Coccolithophores are unicellular phytoplanktonic organisms characterized by a covering of calcite plates, the coccoliths, which are produced intracellularly. These calcifiers, as one of the main planktonic functional groups, play an important role in the inorganic carbon cycle and possibly as ballast that sinks organic carbon to the deep-sea. Most efforts to understanding coccolithophore response to ocean acidification (OA) –or the raise in atmospheric CO2 reduces ocean pH and saturation states (Ω) of CaCO3– have been through lab experiments, mostly using a small set of strains of the cosmopolitan, easily cultivated species Emiliania huxleyi. This species is especially interesting because it is young (~ 291,000 years) and has adapted to a wide range of marine environments. However, it is not the only coccolithophore and even within that species there is a lot of phenotypic and genetic diversity and diverse responses to OA in the lab. Despite the efforts made it is unclear how the physiological effects under controlled conditions translate to community-level responses in the field. This thesis aimed to contribute to understanding this issue by studying the distribution, composition and realized niches of coccolithophore assemblages and E. huxleyi morphotypes in contrasting pCO2/pH/Ωcalcite environments of the Eastern South Pacific, and to evaluate the responses of different E. huxleyi 22 morphotypes to targeted pCO2/pH levels set in the lab. For this, the coccolithophores were surveyed in a coastal-oceanic section, mesotrophic waters, upwelling systems, and fjords- channels of Patagonia. From a total of 40 species, E. huxleyi was the most prevalent (30-100 % relative abundance). Within this taxon, several morphotypes has been described as stable in culture and genetically differentiated (e.g., the A and R morphotypes). The moderately-calcified A morphotype dominated the E. huxleyi populations being only surpassed by the R hyper- calcified morphotype in upwelling systems with high pCO2/low pH. This abrupt shift in the composition of E. huxleyi populations suggested that these coastal environments hold genetic reservoirs for their adaptation to OA. Therefore, the hypothesis was tested that these forms are adapted to resist high pCO2/low pH conditions. Unexpectedly, the morphotypes from the Eastern South Pacific were not more sensitive than the R hyper-calcified strains from neighboring high pCO2/low pH waters (lowering growth rates and PIC/POC ratios). On the other hand, realized-niche analysis showed that the A morphotype has a broader niche that is more tolerant to environmental-change (i.e., generalist) than the R morphotype’s niche, specialized to high pCO2/low pH waters. The lack of evidence for local adaptation to high pCO2/low pH conditions in E. huxleyi, might be explained by a narrow unimodal niche response to Ωcalcite revealed by niche analysis that was not tested experimentally. Alternatively, the R hyper-calcified morphotype might be selected by an unidentified condition particular to the Eastern South Pacific that correlates with temperature, salinity, and Ωcalcite of its realized-niche. Overall, despite their rapid turnover and large population sizes, oceanic planktonic microorganisms do not necessarily exhibit adaptations to high-pCO2 upwelled waters, and this ubiquitous coccolithophore may be near the limit of its capacity to adapt to ongoing OA.