of light. [2] Previous studies have focused mainly on the internal structures of the compound eyes, as they are key to decode the distinguished visual performance of insects. [3][4][5][6][7][8] Some literature data are present on the ultrastructure, material composition, and sclerotization of the corneal cuticle and their influence on insect vision, [9][10][11][12] however, no study has ever established a quantitative link between the structural and biochemical factors and the biomechanical properties of the corneal cuticle. This is especially important, because as a part of insect exoskeleton, eyes should not only possess good optic properties, but also be able to resist mechanical stresses. For instance, the compound eyes should be able to prevent insect head from damage, maintain the mechanical stability between the ommatidia, and support the internal nervous system. [13] Currently, using the existing data, we can hardly explain the mechanisms behind the mechanical stability of corneal cuticle, especially knowing that the elasticity modulus of resilin-rich cuticle (1-60 MPa) is too low to allow for the observed stability. [14] Here, to fill this gap in the literature, we studied the microstructure, sclerotization, and mechanical properties of locust corneal cuticle. The desert locust Schistocerca gregaria was chosen, because it is one of the most established species in studies of cuticle biomechanics. [15][16][17][18][19][20] We used scanning electron microscopy (SEM) and confocal laser scanning microscopy (CLSM) for the structural and material analyses, respectively, and nanoindentation for measuring cuticle stiffness and hardness. Our results enabled us to shed light on the complex relationship between the structure, material, and properties of the corneal cuticle and their influence on the mechanical function of insect eyes.
Results
Microstructure and Material Composition of EyesTransverse fractures on the minor axis of both deep-frozen and air-dried eyes were visualized by SEM. The results revealed that the corneal cuticle consists of the typical layers of epi-, exo-, and endocuticle (Figure 1a,b). The epicuticle is very thin and unstructured, whereas the exocuticle has dense sublayers which are arranged helicoidally, similar to that observed in Compound eyes of insects should be both thin and transparent to allow light to pass through, and at the same time mechanically stable to serve as exoskeleton. These conflicting requirements make the corneal cuticle an interesting example for studying cuticle biomechanics as well as for designing composite materials that seek similar properties. Here, scanning electron microscopy, confocal laser scanning microscopy, and nanoindentation are combined to investigate the microstructure, material composition, and material properties of the corneal cuticle of desert locust Schistocerca gregaria. The results suggest that a fully helicoidal architecture and large proportion of resilin in the corneal cuticle are likely to be adaptations for light transmission. Even though th...