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\title{Environmental and physiological conditions that led to the emergence of
intracellular calcification}



\author[1]{Soenke Hohn}%
\author[2]{Jacopo Dal Corso}%
\author[3]{Kim Hoke}%
\author[4]{Silke Thoms}%
\author[1]{Agostino Merico}%
\affil[1]{Leibniz Centre for Tropical Marine Research}%
\affil[2]{China University of Geosciences State Key Laboratory of Biogeology and Environmental Geology}%
\affil[3]{Colorado State University}%
\affil[4]{Alfred-Wegener-Institut Helmholtz-Zentrum für Polar- und Meeresforschung}%


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  \date{May 27, 2024}


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\begin{abstract}
The emergence of intracellular calcification among marine unicellular
eukaryotes in the Late Triassic (237-201 Ma) had profound consequences
for the carbonate buffering capacity of the ocean. Research on the
appearance of calcification typically focuses on the reasons that made
this process successful on a global scale. The underlying environmental
and physiological conditions that led to its appearance, therefore, are
still obscure. Using gene tree analysis, we show that the physiological
machinery for calcification was already present in non-calcifying
ancestor cells. Additionally, by modelling the energy demands for
calcium transport in calcifying and non-calcifying cells, we demonstrate
that intracellular calcification does not require additional energy
investments. Since all eukaryotic cells export calcium across the plasma
membrane, our findings indicate that the onset of intracellular
calcification in Earth history only required the activation of calcium
transport proteins during their passage to the plasma membrane. Our work
sheds new light on the physiological and biogeochemical conditions that
led to one of the most important evolutionary innovations of the
Mesozoic era.%
\end{abstract}%



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