Mineral homeostasis and regulation of mineralization processes in the skeletons of sharks, rays and relatives (Elasmobranchii)

Mason N. Dean*, Laura Ekstrom, Efrat Monsonego-Ornan, Jim Ballantyne, P. Eckhard Witten, Cyrena Riley, Wouter Habraken, Sidney Omelon

*Corresponding author for this work

Research output: Contribution to journalReview articlepeer-review

38 Scopus citations


Sharks, rays and other elasmobranch fishes are characterized by a skeletal type that is unique among living vertebrates, comprised predominantly of an unmineralized cartilage, covered by a thin outer layer of sub-millimeter, mineralized tiles called tesserae. The mineralized portion of the skeleton appears to grow only by apposition, adding material at the edges of each tessera; maintenance of non-mineralized joints between tesserae is therefore vital, with precise control of mineral deposition and inhibition at the many thousands of growth fronts in the skeleton. Yet, we have only scattered evidence as to how the elasmobranchs mineralize and grow their skeletons. In this review, we take an "environment to skeleton" approach, drawing together research from a vast range of perspectives to track calcium and phosphate from the typical elasmobranch habitats into and through the body, to their deposition at tesseral growth fronts. In the process, we discuss the available evidence for skeletal resorption capability, mineral homeostasis hormones, and nucleation inhibition mechanisms. We also outline relevant theories in crystal nucleation and typical errors in measurements of serum calcium and phosphate in the study of vertebrate biology. We assemble research that suggests consensus in some concepts in elasmobranch skeletal development, but also highlight the very large gaps in our knowledge, particularly in regards to endocrine functional networks and biomineralization mechanisms. In this way, we lay out frameworks for future directions in the study of elasmobranch skeletal biology with stronger and more comparative links to research in other disciplines and into other taxa.

Original languageAmerican English
Pages (from-to)51-67
Number of pages17
JournalSeminars in Cell and Developmental Biology
StatePublished - 1 Oct 2015

Bibliographical note

Funding Information:
This review was inspired to a large degree by Marshall Urist's meticulous study and visionary perspective of elasmobranch skeletal biology. That he is cited in every section of this work is testament to his broad thinking; we wish we had had the chance to meet him. We would like to thank Sergio Bertazzo and John Davey for their invitation and patience during the writing process. Janine Danks, John Dunlop, Gert Flik, Brian Hall, Ron Shahar, Adam Summers, and Byrappa Venkatesh provided support and discussions that shaped this work. Anne Kaulfuß, John Maisey, Ronald Seidel, and Adam Summers generously provided unpublished data and images that greatly enhanced our figures and Pepijn Kamminga shared the CT scan used in Fig. 1 . Kady Lyons and Kazz Kawasaki were invaluable for their help improving specific, clumsy portions of text. MND was supported by an HFSP Young Investigators’ Grant ( RGY0067-2013 ) and DFG-FR 2190/4-1 Gottfried Wilhelm Leibniz-Preis 2010 .

Publisher Copyright:
© 2015 Elsevier Ltd.


  • Apatite
  • Biomineralization
  • Elasmobranch
  • Skeleton
  • Tessellated cartilage
  • Tesserae


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