Living jewels: iterative evolution of iridescent blue leaves from helicoidal cell walls.

BACKGROUND AND AIMS: Structural colour is responsible for the remarkable metallic blue colour seen in the leaves of several plants. Species belonging to only ten genera have been investigated to date, revealing four photonic structures responsible for structurally coloured leaves. One of these is the helicoidal cell wall, known to create structural colour in the leaf cells of five taxa. Here we investigate a broad selection of land plants to understand the phylogenetic distribution of this photonic structure in leaves. METHODS: We identified helicoidal structures in the leaf epidermal cells of 19 species using transmission electron microscopy. Pitch measurements of the helicoids were compared with the reflectance spectra of circularly polarized light from the cells to confirm the structure-colour relationship. RESULTS: By incorporating species examined with a polarizing filter, our results increase the number of taxa with photonic helicoidal cell walls to species belonging to at least 35 genera. These include 19 monocot genera, from the orders Asparagales (Orchidaceae) and Poales (Cyperaceae, Eriocaulaceae, Rapateaceae) and 16 fern genera, from the orders Marattiales (Marattiaceae), Schizaeales (Anemiaceae) and Polypodiales (Blechnaceae, Dryopteridaceae, Lomariopsidaceae, Polypodiaceae, Pteridaceae, Tectariaceae). CONCLUSIONS: Our investigation adds considerably to the recorded diversity of plants with structurally coloured leaves. The iterative evolution of photonic helicoidal walls has resulted in a broad phylogenetic distribution, centred on ferns and monocots. We speculate that the primary function of the helicoidal wall is to provide strength and support, so structural colour could have evolved as a potentially beneficial chance function of this structure.

Structural colours in the frond of Microsorum thailandicum.

Blue and near-ultraviolet structural colours have often been reported in understorey plants living in deep shade. While this intense blue coloration is very catchy to the eye of a human observer, there are cases in which structural colours can be hidden either by the scattered light interacting with pigments or because they are found in unexpected positions in the plants. Here, we show that the fronds of Microsorum thailandicum produce structural coloration on both the adaxial and abaxial epidermal surface. While cellulose helicoidal structures are responsible for this coloration in both epidermal layers, the reflected colours are consistently different: an intense blue reflection is found in the adaxial epidermis while red-shifted and less intense colours are observed in the abaxial epidermis, possibly suggesting photo-adaptation of the plant to the light environment. By comparing the optical properties of the fern with its anatomy we computed the theoretical reflection accounting for the presence of disorder in the cellulose helicoidal architecture.