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Echinoderms and vertebrates both possess mesodermal endoskeletons. In vertebrates, the response to mechanical loads and the capacity to remodel the ultrastructure of the skeletal system are fundamental attributes of their endoskeleton. To determine whether these characteristics are also inherent in Echinoderms, we conducted a comprehensive biomechanical and morphological study on the endoskeleton of Asterias rubens, a representative model organism for Echinoderm skeletons. Our analysis involved high-resolution X-ray CT scans of entire individual ossicles, covering the full stereom distribution along with the attached muscles. Leveraging this data, we conducted finite element analysis to explore the correlation between mechanical loads acting on an ossicle and its corresponding stereom structure. To understand the effects of localized stress concentration, we examined stereom regions subjected to high mechanical stress and compared them to areas with lower mechanical stress. Our results show that the stereom microstructure, both in terms of thickness and orientation, corresponds closely to the mechanical loading experienced by the ossicles. Additionally, by comparing the stereom structures of ossicles in various developmental stages, we assessed the general remodeling capacity of these ossicles. Our findings suggest that the ability to adapt to mechanical loads is a common feature of mesoderm endoskeletons within the Deuterostomia taxonomic group. However, the material remodelling may be a specific trait unique to vertebrate endoskeletons. STATEMENT OF SIGNIFICANCE: This study shows a correlation between the ultrastructure and the mechanical stress in the starfish endoskeleton, suggesting that this fundamental structure-function relationship may be an ancestral feature of not only vertebrate endoskeletons. However, unlike vertebrate skeletons, not all starfish ossicles remodel in response to changing stress, indicating a potential divergence in skeletal adaptation mechanisms. Our methodological approach combines morphometrics and finite element modeling and thus provides a powerful tool to investigate biomechanics in complex skeletal structures., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Ltd.)

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.

The skeletal plates of echinoids consist of a peculiar lightweight structure, called stereom, which is organized in a porous three-dimensional lattice-like meshwork. The stereom is characterized by an extremely complex and diverse microarchitecture, largely varying not only from species to species but also among different test plates. It consists of different basic types combined in extremely different ways according to specific functional needs, creating species-specific structural patterns. These patterns can lead to specific mechanical behaviours, which can inspire biomimetic technology and design development. In this framework, the present study aimed to characterize the species-specific pattern of the Paracentrotus lividus interambulacral plate and the main microstructural features regarding its geometrical variability and mechanical responses. The results achieved quantitatively highlighted the differences between the analysed stereom types providing new insights regarding their topological configuration and isotropic and anisotropic behaviour. Interestingly, data also revealed that the galleried stereom present at the tubercle is significantly different from the one located at the suture. These analyses and findings are encouraging and provide a starting point for future research to unravel the wide range of mechanical strategies evolved in the echinoid skeletal structure.

Repeated polygonal patterns are pervasive in natural forms and structures. These patterns provide inherent structural stability while optimizing strength-per-weight and minimizing construction costs. In echinoids (sea urchins), a visible regularity can be found in the endoskeleton, consisting of a lightweight and resistant micro-trabecular meshwork (stereom). This foam-like structure follows an intrinsic geometrical pattern that has never been investigated. This study aims to analyse and describe it by focusing on the boss of tubercles-spine attachment sites subject to strong mechanical stresses-in the common sea urchin Paracentrotus lividus . The boss microstructure was identified as a Voronoi construction characterized by 82% concordance to the computed Voronoi models, a prevalence of hexagonal polygons, and a regularly organized seed distribution. This pattern is interpreted as an evolutionary solution for the construction of the echinoid skeleton using a lightweight microstructural design that optimizes the trabecular arrangement, maximizes the structural strength and minimizes the metabolic costs of secreting calcitic stereom. Hence, this identification is particularly valuable to improve the understanding of the mechanical function of the stereom as well as to effectively model and reconstruct similar structures in view of future applications in biomimetic technologies and designs.

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)., (© 2020 John Wiley & Sons Ltd.)

Pleurocystitid rhombiferans are among the most unusual echinoderms whose mode of life has been long debated. These echinoderms are usually interpreted as vagile epibenthic echinoderms, moving over the sea bottom by means of a flexible stem. Although their life habits and posture are reasonably well understood, the mechanisms that control the movement of stem are highly controversial. Specifically, it is unknown whether the stem flexibility was under the control of muscles or ligamentary mutable collagenous tissues (MCTs). Here, we reconstruct palaeoanatomy of the two Ordovician pleurocystitid rhombiferans (Pleurocystites and Amecystis) based on stereom microstructure. We show that the articular facets of columnals in pleurocystitid rhombiferans are composed of fine labyrinthic stereom. Comparison with modern echinoderms suggests that this type of stereom was associated with muscles implying that their stem was a muscular locomotory organ supporting an active mode of life.

Growth dynamics of the primary spine of the cidaroid sea urchin Phyllacanthus imperialis was assessed for the first time using pulsed (26) Mg-labeling and NanoSIMS isotopic imaging. The sea urchin was incubated twice (for 48 h) in artificial seawater with elevated level of (26) Mg. After each labeling event, the sea urchin was returned for 72 h to seawater with natural isotopic abundance of (26) Mg. NanoSIMS ion microprobe was subsequently used to visualize the labeled regions of the spine with submicrometer lateral resolution. The growth of the new skeleton was restricted to the distalmost and peripheral portions of the spine. Skeletogenesis involved mostly the deposition of continuous thickening layers and lateral growth involving bridges between previously formed trabeculae. The timescale of formation of individual thickening layers (ca. 1 µm in width) on the stereom trabeculae was on the order of 1 day. Longitudinal growth occurred mainly at the periphery in the form of small portions of the thickening deposits or more massive microspines that appeared to branch and fuse with those above and below. These microspines were found to grow at about 10 µm/day. These results reveal that the skeletal growth of a juvenile cidaroid spine is complex and highly heterogeneous, with different extension rates depending on the stage of the stereom development and/or direction of the growth fronts. The growth pattern observed here at the submicrometer scale provides direct evidence supporting the earlier suggestions that the lamellar structure of echinoderm stereom is formed by periodic deposition of continuous mineral layers., (© 2014 Wiley Periodicals, Inc.)