Protective role of Arapaima gigas fish scales: Structure and mechanical behavior

Wen Yang, Vincent R. Sherman, Bernd Gludovatz, Mason Mackey, Elizabeth A. Zimmermann, Edwin H. Chang, Eric Schaible, Zhao Qin, Markus J. Buehler, Robert O. Ritchie, Marc A. Meyers

Research output: Contribution to journalArticlepeer-review

164 Scopus citations


The scales of the arapaima (Arapaima gigas), one of the largest freshwater fish in the world, can serve as inspiration for the design of flexible dermal armor. Each scale is composed of two layers: a laminate composite of parallel collagen fibrils and a hard, highly mineralized surface layer. We review the structure of the arapaima scales and examine the functions of the different layers, focusing on the mechanical behavior, including tension and penetration of the scales, with and without the highly mineralized outer layer. We show that the fracture of the mineral and the stretching, rotation and delamination of collagen fibrils dissipate a significant amount of energy prior to catastrophic failure, providing high toughness and resistance to penetration by predator teeth. We show that the arapaima's scale has evolved to minimize damage from penetration by predator teeth through a Bouligand-like arrangement of successive layers, each consisting of parallel collagen fibrils with different orientations. This inhibits crack propagation and restricts damage to an area adjoining the penetration. The flexibility of the lamellae is instrumental to the redistribution of the compressive stresses in the underlying tissue, decreasing the severity of the concentrated load produced by the action of a tooth. The experimental results, combined with small-angle X-ray scattering characterization and molecular dynamics simulations, provide a complete picture of the mechanisms of deformation, delamination and rotation of the lamellae during tensile extension of the scale.

Original languageEnglish (US)
Pages (from-to)3599-3614
Number of pages16
JournalActa Biomaterialia
Issue number8
StatePublished - Aug 2014
Externally publishedYes


  • Arapaima
  • Armor
  • Collagen
  • Delamination
  • Fish scales

ASJC Scopus subject areas

  • Biotechnology
  • Biomaterials
  • Biochemistry
  • Biomedical Engineering
  • Molecular Biology


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