Optical properties of the jellyfish surface above the waterline: microvillar array in pleustonic hydrozoans

The transparent jellyfish body is often difficult to see underwater, as its refractive index is similar to that of seawater, resulting in a low light reflectance on the body surface. Nevertheless, the outlines of jellyfish can be recognized by the slight reflection of light from their body surfaces. In some jellyfish species, the epidermis covering the body surface has an array of microvilli, nanostructures that can potentially reduce light reflection. However, the anti-reflective effect is minimal in water, as the difference in the refractive indices of tissue and seawater is so small that reflectance is low, even on flat surfaces. In jellyfish that have pneumatophores, structures used in floating and drifting on the sea surface, light reflection on the surface is expected to be large and noticeable owing to the large differences in refractive indices between the pneumatophore exposed above the water surface and air. In the current study, we examined the epidermal ultrastructure and refractive index of the pneumatophores of a Portuguese man o' war (Physalia physalis) and a by-the-wind sailor (Velella velella).

The refractive index of P. physalis pneumatophores measured with an Abbe refractometer was approximately 1.344. Microvillar arrays were found in epidermal cells of both P. physalis and V. velella. Based on the length, thickness, and pitch of the microvilli, we constructed simplified structural models for the simulation of light reflection using rigorous coupled wave analysis (RCWA). Our simulations showed that reflectance on the microvillar models could be greater or less than that on the flat surface, depending on light conditions (wavelength and angle of incidence), but with an overall effect of reduced reflection. Reflection reduction in microvillar models was particularly significant at large incident angles, where reflectance was extremely high on the flat surface.

Microvillar arrays found on the epidermis potentially reduce surface reflections of the pneumatophore and contribute to the reduction in visibility of the pleustonic hydrozoans above the sea surface. Moreover, less reflection at the pneumatophore surface indicates greater transmission of light through transparent bodies, potentially providing a counter-illumination effect that obscures the shadow of the hydrozoan bodies, depending on the intensity of ambient light.