A new structural model of bivalve ligament from Solen grandis

It is widely accepted that the structure of bivalve ligament contains two layers: outer organic lamellar and inner aragonite fibrous layer. However, by using scanning electron microscopy (SEM), X-ray diffraction (XRD), and infrared spectroscopy (FTIR), we found an additional protein fibrous layer in the Solen grandis ligament. The ligament consists of three layers: an outer layer (OL), middle layer (ML), and inner layer (IL). OL consists of lamellar protein, oriented normal to dorsal edge. ML has unique spring-like protein fibers (ca. 120 nm in diameter) structure, stretching continuously from the left to right valve. Interestingly, IL comprises of aragonite fibers (ca. 100 nm in diameter) oriented posteriorly and inclined at an angle about 26° to dorsal edge at dorsal part; then they turn gradually to ventral part, showing striking zigzag structure vertical to ventral surface (VS). The ligament presents a new three-layered structural model different from previous reports; it may inspire not only in materials synthesis and mechanical design, but also in bivalve classification and evolution.