@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 {Helliwell2020,
	title = {A novel single-domain Na +-selective voltage-gated channel in photosynthetic eukaryotes},
	journal = {bioRxiv},
	year = {2020},
	note = {Publisher: Cold Spring Harbor Laboratory tex.mendeley-tags: RCC1456},
	month = {apr},
	pages = {2020.04.29.068528},
	abstract = {The evolution of Na+-selective four-domain voltage-gated channels (4D-Navs) in animals allowed rapid Na+-dependent electrical excitability, and enabled the development of sophisticated systems for rapid and long-range signalling. Whilst bacteria encode single-domain Na+-selective voltage-gated channels (BacNav), they typically exhibit much slower kinetics than 4D-Navs, and are not thought to have crossed the prokaryote-eukaryote boundary. As such, the capacity for rapid Na+-selective signalling is considered to be confined to certain animal taxa, and absent from photosynthetic eukaryotes. Certainly, in land plants, such as the Venus Flytrap where fast electrical excitability has been described, this is most likely based on fast anion channels. Here, we report a unique class of eukaryotic Na+-selective single-domain channels (EukCatBs) that are present primarily in haptophyte algae, including the ecologically important calcifying coccolithophores. The EukCatB channels exhibit very rapid voltage-dependent activation and inactivation kinetics, and sensitivity to the highly selective 4D-Nav blocker tetrodotoxin. The results demonstrate that the capacity for rapid Na+-based signalling in eukaryotes is not restricted to animals or to the presence of 4D-Navs. The EukCatB channels therefore represent an independent evolution of fast Na+-based electrical signalling in eukaryotes that likely contribute to sophisticated cellular control mechanisms operating on very short time scales in unicellular algae. One Sentence Summary The capacity for rapid Na+-based signalling has evolved in ecologically important coccolithophore species via a novel class of voltage-gated Na+ channels, EukCatBs.},
	keywords = {RCC1456},
	doi = {10.1101/2020.04.29.068528},
	url = {https://doi.org/10.1101/2020.04.29.068528},
	author = {Helliwell, Katherine E and Chrachri, Abdul and Koester, Julie and Wharam, Susan and Wheeler, Glen L and Brownlee, Colin}
}
@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 {Durak2016,
	title = {A role for diatom-like silicon transporters in calcifying coccolithophores},
	journal = {Nature Communications},
	volume = {7},
	number = {February},
	year = {2016},
	note = {tex.mendeley-tags: 2016,RCC1130,RCC1303,RCC1453,RCC1456,RCC3432},
	pages = {10543},
	abstract = {Biomineralisation by marine phytoplankton, such as the silicifying diatoms and calcifying coccolithophores, plays an important role in carbon and nutrient cycling in the oceans. Silicification and calcification are distinct cellular processes with no known common mechanisms. As a result, it is thought that coccolithophores are able to outcompete diatoms in Si-depleted waters, which can contribute to the formation of coccolithophore blooms. Here, we show that an expanded family of diatom-like silicon transporters (SITs) are present in both silicifying and calcifying haptophyte phytoplankton, including some coccolithophores of global ecological importance. We find an essential role for Si in calcification in these coccolithophores, indicating that Si uptake contributes to the very different forms of biomineralisation in diatoms and coccolithophores. However, SITs and the requirement for Si are significantly absent from the highly abundant bloom-forming coccolithophores, such as Emiliania huxleyi. These very different requirements for Si in coccolithophores are likely to have major influence on their competitive interactions with diatoms and other siliceous phytoplankton.},
	keywords = {(RCC1130, (RCC1456), 2016, biomineralisation, coccolithophores, Gephyrocapsa oceanica (RCC1303) and Scyphosphaera, haptophytes, RCC1130, RCC1303, RCC1453, RCC1456, RCC3432, silica, TMR5 (RCC3432{\textemdash}Sea of Japan) and PZ241 (RCC1453{\textemdash}Med},
	issn = {2041-1723},
	doi = {10.1038/ncomms10543},
	author = {Durak, Grazyna M and Taylor, Alison R and Probert, Ian and de Vargas, Colomban and Audic, St{\'e}phane and Schroeder, Declan C and Brownlee, Colin and Wheeler, Glen L}
}
@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}
}
