It has been argued that concentration of major metallic ions such as Mg 2+ and Ca 2+ plays a role in determining the composition of the echinoderm skeleton. Consequently, in several studies Mg/Ca ratio from modern and fossil echinoderm ossicles was used as a reliable proxy of secular Mg 2+ /Ca 2+ changes of Phanerozoic seawater. However, although significant progress has been made in understanding biomineralization of echinoderms, it is still largely unknown what are the sources and physiological pathways of major ions that contribute to skeleton formation. Herein we tested the effects of modifications of ambient seawater Mg 2+ /Ca 2+ ratio (~5) and Mg-enrichment of the diet on the Mg/Ca ratio in regenerating spines of sea urchin Paracentrotus lividus under experimental conditions. We found that sea urchins cultured in seawater with decreased Mg 2+ /Ca 2+ ratio (~1.9) produced a skeleton with also decreased Mg/Ca ratio. However, the skeleton of specimens fed on a Mg-enriched diet showed significantly higher Mg/Ca ratio. This suggests that the seawater is an important but not the only source of ions that contributes to the Mg/Ca ratio of the skeleton. Consequently, the reliability of geochemical models that link directly seawater chemistry with the Mg/Ca ratio of the skeleton should be reevaluated.
It has been argued that skeletal Mg/Ca ratio in echinoderms is mostly governed by Mg 2+ and Ca 2+ concentrations in the ambient seawater. Accordingly, well-preserved fossil echinoderms were used to reconstruct Phanerozoic seawater Mg 2+ /Ca 2+ ratio. However, Mg/Ca ratio in echinoderm skeleton can be affected by a number of environmental and physiological factors, the effects of which are still poorly understood. Notably, experimental data supporting the applicability of echinoderms in paleoenvironmental reconstructions remain limited. Here, we investigated the effect of ambient Mg 2+ /Ca 2+ seawater ratio and diet on skeletal Mg/Ca ratio and growth rate in two echinoid species (Psammechinus miliaris and Prionocidaris baculosa). Sea urchins were tagged with manganese and then cultured in different Mg 2+ /Ca 2+ conditions to simulate fluctuations in the Mg 2+ /Ca 2+ seawater ratios in the Phanerozoic. Simultaneously, they were fed on a diet containing different amounts of magnesium. Our results show that the skeletal Mg/Ca ratio in both species varied not only between ossicle types but also between different types of stereom within a single ossicle. Importantly, the skeletal Mg/Ca ratio in both species decreased proportionally with decreasing seawater Mg 2+ /Ca 2+ ratio. However, sea urchins feeding on Mg-enriched diet produced a skeleton with a higher Mg/Ca ratio. We also found that although incubation in lower ambient Mg 2+ /Ca 2+ ratio did not affect echinoid respiration rates, it led to a decrease or inhibition of their growth. Overall, these results demonstrate that although skeletal Mg/Ca ratios in echinoderms can be largely determined by seawater chemistry, the type of diet may also influence skeletal geochemistry, which imposes constraints on the application of fossil echinoderms as a reliable proxy. The accuracy of paleoseawater Mg 2+ /Ca 2+ calculations is further limited by the fact that Mg partition coefficients vary significantly at different scales (between species, specimens feeding on different types of food, different ossicle types, and stereom types within a single ossicle).
Owing to their remarkable physical properties, cellular structures, such as triply periodic minimal surfaces (TPMS), have multidisciplinary and multifunctional applications. Although these structures are observed in nature, examples of TPMS with large length scales in living organisms are exceedingly rare. Recently, microstructure reminiscent of the diamond-type TPMS was documented in the skeleton of the modern knobby starfish Protoreaster nodosus . Here we report a similar microlattice in a 385 Myr old crinoid Haplocrinites , which pushes back the origins of this highly ordered microstructure in echinoderms into the Devonian. Despite the low Mg 2+ /Ca 2+ ratio of the ‘calcite’ Devonian sea, the skeleton of these crinoids has high-Mg content, which indicates strong biological control over biomineralogy. We suggest that such an optimization of trabecular arrangement additionally enriched in magnesium, which enhances the mechanical properties, might have evolved in these crinoids in response to increased predation pressure during the Middle Palaeozoic Marine Revolution. This discovery illustrates the remarkable ability of echinoderms, through the process of evolutionary optimization, to form a lightweight, stiff and damage-tolerant skeleton, which serves as an inspiration for biomimetic materials.
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