@article {vazquez_high-co2_2022,
	title = {High-CO2 Levels Rather than Acidification Restrict Emiliania huxleyi Growth and Performance},
	journal = {Microbial Ecology},
	year = {2022},
	abstract = {The coccolithophore Emiliania huxleyi shows a variety of responses to ocean acidification (OA) and to high-CO2 concentrations, but there is still controversy on differentiating between these two factors when using different strains and culture methods. A heavily calcified type A strain isolated from the Norwegian Sea was selected and batch cultured in order to understand whether acclimation to OA was mediated mainly by CO2 or H+, and how it impacted cell growth performance, calcification, and physiological stress management. Emiliania huxleyi responded differently to each acidification method. CO2-enriched aeration (1200 {\textmu}atm, pH 7.62) induced a negative effect on the cells when compared to acidification caused by decreasing pH alone (pH 7.60). The growth rates of the coccolithophore were more negatively affected by high pCO2 than by low pH without CO2 enrichment with respect to the control (400 {\textmu}atm, pH 8.1). High CO2 also affected cell viability and promoted the accumulation of reactive oxygen species (ROS), which was not observed under low pH. This suggests a possible metabolic imbalance induced by high CO2 alone. In contrast, the affinity for carbon uptake was negatively affected by both low pH and high CO2. Photochemistry was only marginally affected by either acidification method when analysed by PAM fluorometry. The POC and PIC cellular quotas and the PIC:POC ratio shifted along the different phases of the cultures; consequently, calcification did not follow the same pattern observed in cell stress and growth performance. Specifically, acidification by HCl addition caused a higher proportion of severely deformed coccoliths, than CO2 enrichment. These results highlight the capacity of CO2 rather than acidification itself to generate metabolic stress, not reducing calcification.},
	keywords = {Calcification, coccolithophores, Emiliania huxleyi, Ocean acidification, pCO2, Photochemistry, phytoplankton, rcc1226, Stress},
	issn = {1432-184X},
	doi = {10.1007/s00248-022-02035-3},
	url = {https://doi.org/10.1007/s00248-022-02035-3},
	author = {V{\'a}zquez, V{\'\i}ctor and Le{\'o}n, Pablo and Gordillo, Francisco J. L. and Jim{\'e}nez, Carlos and Concepci{\'o}n, I{\~n}iguez and Mackenzie, Kevin and Bresnan, Eileen and Segovia, Mar{\'\i}a}
}
@article {langer_role_2021,
	title = {Role of silicon in the development of complex crystal shapes in coccolithophores},
	journal = {New Phytologist},
	volume = {231},
	number = {5},
	year = {2021},
	note = {_eprint: https://onlinelibrary.wiley.com/doi/pdf/10.1111/nph.17230},
	pages = {1845{\textendash}1857},
	abstract = {The development of calcification by the coccolithophores had a profound impact on ocean carbon cycling, but the evolutionary steps leading to the formation of these complex biomineralized structures are not clear. Heterococcoliths consisting of intricately shaped calcite crystals are formed intracellularly by the diploid life cycle phase. Holococcoliths consisting of simple rhombic crystals can be produced by the haploid life cycle stage but are thought to be formed extracellularly, representing an independent evolutionary origin of calcification. We use advanced microscopy techniques to determine the nature of coccolith formation and complex crystal formation in coccolithophore life cycle stages. We find that holococcoliths are formed in intracellular compartments in a similar manner to heterococcoliths. However, we show that silicon is not required for holococcolith formation and that the requirement for silicon in certain coccolithophore species relates specifically to the process of crystal morphogenesis in heterococcoliths. We therefore propose an evolutionary scheme in which the lower complexity holococcoliths represent an ancestral form of calcification in coccolithophores. The subsequent recruitment of a silicon-dependent mechanism for crystal morphogenesis in the diploid life cycle stage led to the emergence of the intricately shaped heterococcoliths, enabling the formation of the elaborate coccospheres that underpin the ecological success of coccolithophores.},
	keywords = {biomineralization, Calcification, coccolith, coccolithophore, evolution, rcc, RCC1178, RCC1181, RCC1456, RCC1460, RCC1461, RCC1477, RCC1800, RCC1801, RCC3777, RCC6506, silicon},
	issn = {1469-8137},
	doi = {10.1111/nph.17230},
	url = {https://onlinelibrary.wiley.com/doi/abs/10.1111/nph.17230},
	author = {Langer, Gerald and Taylor, Alison R. and Walker, Charlotte E. and Meyer, Erin M. and Ben Joseph, Oz and Gal, Assaf and Harper, Glenn M. and Probert, Ian and Brownlee, Colin and Wheeler, Glen L.}
}
@article {gebuhr_strain-specific_2021,
	title = {Strain-specific morphological response of the dominant calcifying phytoplankton species Emiliania huxleyi to salinity change},
	journal = {PLOS ONE},
	volume = {16},
	number = {2},
	year = {2021},
	note = {Publisher: Public Library of Science},
	month = {feb},
	pages = {e0246745},
	abstract = {The future physiology of marine phytoplankton will be impacted by a range of changes in global ocean conditions, including salinity regimes that vary spatially and on a range of short- to geological timescales. Coccolithophores have global ecological and biogeochemical significance as the most important calcifying marine phytoplankton group. Previous research has shown that the morphology of their exoskeletal calcified plates (coccoliths) responds to changing salinity in the most abundant coccolithophore species, Emiliania huxleyi. However, the extent to which these responses may be strain-specific is not well established. Here we investigated the growth response of six strains of E. huxleyi under low (ca. 25) and high (ca. 45) salinity batch culture conditions and found substantial variability in the magnitude and direction of response to salinity change across strains. Growth rates declined under low and high salinity conditions in four of the six strains but increased under both low and high salinity in strain RCC1232 and were higher under low salinity and lower under high salinity in strain PLYB11. When detailed changes in coccolith and coccosphere size were quantified in two of these strains that were isolated from contrasting salinity regimes (coastal Norwegian low salinity of ca. 30 and Mediterranean high salinity of ca. 37), the Norwegian strain showed an average 26\% larger mean coccolith size at high salinities compared to low salinities. In contrast, coccolith size in the Mediterranean strain showed a smaller size trend (11\% increase) but severely impeded coccolith formation in the low salinity treatment. Coccosphere size similarly increased with salinity in the Norwegian strain but this trend was not observed in the Mediterranean strain. Coccolith size changes with salinity compiled for other strains also show variability, strongly suggesting that the effect of salinity change on coccolithophore morphology is likely to be strain specific. We propose that physiological adaptation to local conditions, in particular strategies for plasticity under stress, has an important role in determining ecotype responses to salinity.},
	keywords = {Calcification, Carbonates, Ecophysiology, Marine ecology, Marine geology, Morphometry, Paleoclimatology, rcc1210, RCC1232, rcc1824, rcc868, RCC904, salinity},
	issn = {1932-6203},
	doi = {10.1371/journal.pone.0246745},
	url = {https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0246745},
	author = {Geb{\"u}hr, Christina and Sheward, Rosie M. and Herrle, Jens O. and Bollmann, J{\"o}rg}
}
@article {Meyer2020,
	title = {Sr in coccoliths of Scyphosphaera apsteinii: Partitioning behavior and role in coccolith morphogenesis},
	journal = {Geochimica et Cosmochimica Acta},
	volume = {285},
	year = {2020},
	note = {Publisher: Elsevier Ltd tex.mendeley-tags: RCC1456},
	month = {sep},
	pages = {41{\textendash}54},
	abstract = {Coccolithophores are important contributors to global calcium carbonate through their species-specific production of calcite coccoliths. Nannofossil coccolith calcite remains an important tool for paleoreconstructions through geochemical analysis of isotopic and trace element incorporation including Sr, which is a potential indicator of past surface ocean temperature and productivity. Scyphosphaera apsteinii (Zygodiscales) exhibits an unusually high Sr/Ca ratio and correspondingly high partitioning coefficient (DSr = 2.5) in their two morphologically distinct types of coccoliths: flat muroliths and barrel-like lopadoliths. Whether or not this reflects mechanistic differences in calcification compared to other coccolithophores is unknown. We therefore examined the possible role of Sr in S. apsteinii calcification by growing cells in deplete (0.33 mmol/mol Sr/Ca), ambient (9 mmol/mol Sr/Ca), and higher than ambient Sr conditions (36 and 72 mmol/mol Sr/Ca). The effects on growth, quantum efficiency of photosystem II (Fv/Fm), coccolith morphology, and calcite DSr were evaluated. No effect on S. apsteinii growth rate or Fv/Fm was observed when cells were grown in Sr/Ca between 0.33{\textendash}36 mmol/mol. However, at 72 mmol/mol Sr/Ca growth rate was significantly reduced, although Fv/Fm was unaffected. Reducing the Sr/Ca from ambient (9 mmol/mol) did not significantly alter the frequency of malformed and aberrant muroliths and lopadoliths, but at higher than ambient Sr/Ca conditions coccolith morphology was significantly disrupted. This implies that Sr is not a critical determining factor in normal coccolith calcite morphology in this dimorphic species. Using energy dispersive spectroscopy (EDS) we observed an increase in [Sr] and decrease in DSr of coccoliths as the Sr/Ca of the growth medium increased. Interestingly, muroliths had significantly lower Sr/Ca than lopadoliths at ambient and elevated [Sr], and lopadolith tips had lower Sr than bases in ambient conditions. In summary, the Sr fractionation behavior of S. apsteinii is distinct from other coccolithophores because of an unusually high DSr and inter- and intra-coccolith variability in Sr/Ca. These observations could be explained by mechanistic differences in the selectivity of the Ca2+ transport pathway or in the Sr-and Ca-binding capacity of organic components, such as polysaccharides associated with coccolithogenesis.},
	keywords = {biomineralization, Calcification, coccolith, coccolithophore, Energy dispersive spectroscopy, Fractionation, RCC1456, Sr/Ca, Strontium, Trace element},
	issn = {00167037},
	doi = {10.1016/j.gca.2020.06.023},
	author = {Meyer, Erin M. and Langer, Gerald and Brownlee, Colin and Wheeler, Glen L. and Taylor, Alison R.}
}
@article {Fox2018,
	title = {Calcein staining as a tool to investigate coccolithophore calcification},
	journal = {Frontiers in Marine Science},
	volume = {5},
	number = {September},
	year = {2018},
	note = {tex.mendeley-tags: RCC 1461,RCC 3777,RCC1130,RCC1303,RCC1456},
	abstract = {This brief paper summarizes the literature on academic accommodations for students with psychiatric disabilities. A definition of psychiatric disability precedes a brief summary of the following specific psychiatric diagnoses: depression, bipolar affective disorder; borderline personality disorder; schizophrenia; and anxiety disorders. Also noted are behavior or personality disorders specifically excluded from coverage under the Americans with Disabilities Act. Functional limitations of this population that may affect academic performance are then listed and defined. Among these are medication side effects, sustaining concentration, maintaining stamina, interacting with others, responding to negative feedback, responding to change, and severe test anxiety. Examples of appropriate instructional strategies are followed by a discussion of reasonable accommodations to provide these students with equal access to the curriculum. Examples are then given of classroom accommodations, examination accommodations, and assignment accommodations. Characteristics of accommodations that are not reasonable are also listed. The paper concludes with a resource list including Web site and contact information for the DO-IT (Disabilities, Opportunities, Internetworking, and Technology) Project. (DB)},
	keywords = {bet hedging, calcein, Calcification, coccolith, coccolithophore, flow cytometry, haptophyte, RCC1130, RCC1303, RCC1456, RCC1461, RCC3777, secretion},
	issn = {2296-7745},
	doi = {10.3389/fmars.2018.00326},
	url = {https://www.frontiersin.org/article/10.3389/fmars.2018.00326/full},
	author = {Fox, Emily and Meyer, Erin and Panasiak, Natalie and Taylor, Alison R.}
}
@article {Walker2018,
	title = {The requirement for calcification differs between ecologically important coccolithophore species},
	journal = {New Phytologist},
	volume = {in press},
	year = {2018},
	note = {tex.mendeley-tags: RCC1731},
	month = {jun},
	keywords = {Calcification, coccolithophore, Coccolithus braarudii, Emiliania huxleyi, phytoplankton, rcc1731},
	issn = {0028646X},
	doi = {10.1111/nph.15272},
	url = {http://doi.wiley.com/10.1111/nph.15272},
	author = {Walker, Charlotte E. and Taylor, Alison R. and Langer, Gerald and Durak, Gra{\.z}yna M. and Heath, Sarah and Probert, Ian and Tyrrell, Toby and Brownlee, Colin and Wheeler, Glen L.}
}
@article {Gerecht2015,
	title = {Phosphorus availability modifies carbon production in Coccolithus pelagicus (Haptophyta)},
	journal = {Journal of Experimental Marine Biology and Ecology},
	volume = {472},
	year = {2015},
	note = {tex.mendeley-tags: RCC1200},
	month = {nov},
	pages = {24{\textendash}31},
	abstract = {The coccolithophore Coccolithus pelagicus (Wallich) Schiller fixes CO{\textexclamdown}inf{\textquestiondown}2{\textexclamdown}/inf{\textquestiondown} into particulate organic carbon (POC) through photosynthesis and into particulate inorganic carbon (PIC) in the form of calcite. To examine the role of phosphorus (P) availability in the production of POC and PIC, C. pelagicus subsp. braarudii (Gaarder) Geisen et al. was grown in semi-continuous cultures at three initial phosphate concentrations (P-replete, 1, and 0.5??M [P]). Reduced P-availability (1 and 0.5??M [P]) decreased POC production, while PIC production only decreased when phosphate concentrations became growth limiting (0.5??M [P]). This decrease has not been observed previously in batch cultures, highlighting the inadequacy of the batch culture approach with regard to determining carbon production. The reduction in growth rate by 50\% at 0.5??M [P] was accompanied by a doubling in cell volume (and POC). PIC production was halved, resulting in a lowered PIC to POC ratio. The average number of coccoliths per cell (and PIC content) remained the same among treatments, despite the significant change in cell size. Our data suggest that POC production in C. pelagicus is more sensitive towards a moderate reduction in phosphorus availability than PIC production. Once phosphorus availability limits cell division, however, phosphorus resources are invested into POC rather than PIC production. This reduces cell density and sinking rates, indicating that coccoliths do not act as ballast for reaching deeper nutrient-rich layers under nutrient limitation.},
	keywords = {Calcification, Carbon production, Coccolithus pelagicus, Phosphorus limitation, RCC1200, Semi-continuous culture},
	issn = {00220981},
	doi = {10.1016/j.jembe.2015.06.019},
	url = {http://linkinghub.elsevier.com/retrieve/pii/S0022098115001756},
	author = {Gerecht, A.C. and {\v S}upraha, L. and Edvardsen, B. and Langer, G. and Henderiks, J.}
}
@article {Drescher2012,
	title = {Coccolithogenesis in scyphosphaera apsteinii (prymnesiophyceae)},
	journal = {Journal of Phycology},
	volume = {48},
	number = {6},
	year = {2012},
	note = {tex.mendeley-tags: RCC1456},
	pages = {1343{\textendash}1361},
	abstract = {Coccolithophores are the most significant producers of marine biogenic calcite, although the intracellular calcification process is poorly understood. In the case of Scyphosphaera apsteinii Lohmann 1902, flat ovoid muroliths and bulky, vase-shaped lopadoliths with a range of intermediate morphologies may be produced by a single cell. This polymorphic species is within the Zygodiscales, a group that remains understudied with respect to ultrastructure and coccolith ontogeny. We therefore undertook an analysis of cell ultrastructure, morphology, and coccolithogenesis. The cell ultrastructure showed many typical haptophyte features, with calcification following a similar pattern to that described for other heterococcolith bearing species including Emiliania huxleyi. Of particular significance was the reticular body role in governing fine-scale morphology, specifically the central pore formation of the coccolith. Our observations also highlighted the essential role of the inter- and intracrystalline organic matrix in growth and arrangement of the coccolith calcite. S. apsteinii secreted mature coccoliths that attached to the plasma membrane via fibrillar material. Time-lapse light microscopy demonstrated secretion of lopadoliths occurred base first before being actively repositioned at the cell surface. Significantly, growth irradiance influenced the coccosphere composition with fewer lopadoliths being formed relative to muroliths at higher light intensities. Overall, our observations support dynamic metabolic (i.e., in response to growth irradiance), sensory and cytoskeletal control over the morphology and secretion of polymorphic heterococcoliths. With a basic understanding of calcification established, S. apsteinii could be a valuable model to further study coccolithophore calcification and cell physiological responses to ocean acidification.},
	keywords = {Calcification, coccolithogenesis, coccolithophore, organic matrix, rcc, RCC1456, reticular body, secretion, ultrastructure, Zygodiscales},
	doi = {10.1111/j.1529-8817.2012.01227.x},
	url = {http://dx.doi.org/10.1111/j.1529-8817.2012.01227.x},
	author = {Drescher, Brandon and Dillaman, Richard M and Taylor, Alison R}
}
@article {Langer2011a,
	title = {CO2 mediation of adverse effects of seawater acidification in Calcidiscus leptoporus},
	journal = {Geochemistry Geophysics Geosystems},
	volume = {12},
	number = {5},
	year = {2011},
	note = {ISBN: 1525-2027 tex.mendeley-tags: 2011,RCC1135,rcc},
	pages = {1{\textendash}8},
	abstract = {The coccolithophore Calcidiscus leptoporus (strain RCC1135) was grown in dilute batch culture at CO2 levels ranging from \~200 to \~1600 matm. Increasing CO2 concentration led to an increased percentage of malformed coccoliths and eventually (at \~1500 matm CO2) to aggregation of cells. Carbonate chemistry of natural seawater was manipulated in three ways: first, addition of acid; second, addition of a HCO3 -/CO3 2- solution; and third, addition of both acid and HCO3 -/CO32- solution. The data set allowed the disentangling of putative effects of the different parameters of the carbonate system. It is concluded that CO2 is the parameter of the carbonate system which causes both aberrant coccolithogenesis and aggregation of cells.},
	keywords = {2011, Calcification, coccolithophores, morphology, ocean acidification., rcc, RCC1135},
	issn = {1525-2027},
	doi = {10.1029/2010GC003393},
	url = {http://www.agu.org/pubs/crossref/2011/2010GC003393.shtml},
	author = {Langer, Gerald and Bode, Maya}
}
