Material and structural properties of fin whale (Balaenoptera physalus) Zwischensubstanz


Correspondence to: Robert E. Shadwick; Department of Zoology, University of British Columbia, 6270 University Blvd., Vancouver, British Columbia V6T 1Z4, Canada. E-mail:


The oral anatomy of the fin whale (Balaenoptera physalus) consists of several major structures crucial to its engulfment method of feeding, such as stiff keratinized baleen plates, a large flaccid tongue, and a prominent vomer. One under-documented part of this anatomy is the cream white Zwischensubstanz that holds the baleen plates to the rostrum at their dorsal base. The mechanical and structural properties of Zwischensubstanz play a key role in baleen plate dynamics and, on the grand scale, contribute to baleen whales' filtration efficiency and attainment of large body size. Compression and tensile tests on the Zwischensubstanz sampled from an 18 m fin whale showed that this material unexpectedly exhibits linear isotropic behaviour with Elastic Modulus of 2.56 ± 0.60 MPa and hysteresis of 0.44 ± 0.02 in compression despite apparent unidirectional growth. Acting similar to a soft rubber, the Zwischensubstanz absorbs and dissipates the enormous forces acting on baleen plates during engulfment feeding while maintaining spacing between the plates to maximize filtration efficiency. Microscopic analysis provided images of connective tissue papillae penetrating the base of the Zwischensubstanz and developing within it to emerge as fully formed, keratinized baleen plates. The plates develop from the papillae and a connective tissue sheet within the 5–7 cm deep Zwischensubstanz. The Zwischensubstanz provides a keratin matrix of concentrically oriented fibers around each papilla forming the hard baleen plates and frayed fringes used for filter feeding. During this formation, the Zwischensubstanz remains unchanged and appears to slough away to allow the baleen plate to grow unhindered. J. Morphol. 274:947–955, 2013. © 2013 Wiley Periodicals, Inc.