Strength in Weakness: The Mutable Collagenous Tissue of Echinoderms

Mutable collagenous tissue (MCT) is a type of connective tissue that is characterized by its capacity to undergo rapid, nervously mediated changes in mechanical properties. In terms of both the magnitude of these changes and the timescale within which they occur (less than one second to a few minutes), this tissue appears to be unique to the phylum Echinodermata and, as it is ubiquitous in all five extant echinoderm classes, it represents one of the four major defining features of the phylum, together with pentaradial symmetry, endoskeletal stereom (calcite meshwork), and the water vascular system. MCT has been the subject of intensive scientific investigation for over 50 years. The primary aim of this contribution is to provide a comprehensive and definitive survey of the current state of knowledge of this remarkable tissue. After outlining the history of the scientific investigation of MCT, we review current information on its anatomical distribution, organization at the histological, ultrastructural and molecular levels, and physiology—focusing on its mechanical behavior and the regulation of this behavior; its significance for echinoderm biology, including pathology; and biomedical and other applications that exploit MCT-derived components or biological principles. We conclude by drawing attention to more serious deficiencies in the current knowledge base and suggesting how these should be rectified.