Priemel, T. | bioRxiv Science

bioRxiv2026

Intracellular photonic crystals in photosynthetic sea slugs form via a kidney-mediated biomineralisation pathway

Sea slugs in the Sacoglossa superorder are some of the few animals capable of photosynthesising by isolating and maintaining functional chloroplasts within their body1,2. While this ability allows some species in this superorder, such as Elysia viridis, to appear green, camouflaging themselves within their surroundings3,4, this species is marked by extremely bright, coloured regions. Here, we show that these animals produce a yet undiscovered class of photonic structure consisting of intracellular mixed amorphous CaCO3 and calcite spherical nanoparticles organised in non-closed-packed face-centred cubic (FCC) lattices and photonic glasses5. By mapping the distribution of the cells containing such architectures, we suggest that their colour is linked both to their function and to their biological formation via the animals renal system. Using a combination of different optical methods and cryo-electron microscopy, we reveal that the biomineralisation pathway proceeds through stages of calcium ion concentration in the kidney, transport via internal vessels, and precipitation from a dense liquid-like precursor, culminating in the formation of monodisperse nanoparticles, which are the building blocks of these photonic structures.

biophysics↗

bioRxiv2025

Nudibranch color diversity shares a common origin in guanine photonic structures

Nudibranchs are well known for their bright and diverse color patterns. This coloration is typically a form of aposematism, warning predators against toxic compounds sequestered from their prey and weaponized as a form of defense. Although many of the hues in nudibranchs have pigmentary origin, multilayer structures composed of guanine nano-platelets have been suggested as the source of color enhancement in the nudibranch Flabellina iodenea. Here, using a combination of white light and Raman microspectroscopy techniques, we report that such guanine-based multilayer structures are a widespread mechanism to create angular-independent structural color across the dorid and aeolid groups. Additionally, by using cryo-FIB tomography, we were able to access the complex 3D organization of the guanine nano-platelets responsible for the strong blue coloration of Chromodoris annae. We propose that the multilayer organization of guanine platelets with varying orientations across the tissue offers a particularly effective strategy for producing diverse optical effects. In this configuration, hue is mainly governed by interlayer spacing, while the angular dependence of color can be tuned through the degree of local order, allowing a single structural motif to generate a broad palette of optical appearances. Significance statementNudibranchs are an extraordinarily diverse group of marine animals, renowned for their dazzling range of colors and striking patterns. Whilst their pigmentary coloration is well understood, so far, structural coloration, obtained only by nanostructures, has only been reported in the nudibranch Flabellina iodenea. In this work, we present a comparative analysis of structural coloration across nudibranch species from both benthic and coral reef environments, and we show that guanine-based nano-structures are a common motif responsible for a wide range of colors, spanning the dorid and aeolid groups. We foresee that the 3D imaging conducted here may serve as inspiration for bio-photonics studies in other marine organisms, and that the structures themselves could serve as inspiration for bio-inspired materials.

biophysics↗