Four hundred years of cork imaging: New advances in the characterization of the cork structure.

In 1665, Robert Hooke was the first to observe cork cells and their characteristic hexagonal shape, using the first optical microscope, which was invented by him at that time. With the evolution of imaging techniques, the structure of cork has been analysed with greater accuracy over time. This work presents the latest advances in the characterization of this unique material through a multiscale approach. Such investigation brings new insight into the architecture of cork, particularly the differences between the cells of the phellem and those bordering the lenticels. In the latter case, cell differentiation from the lenticular phellogen was restricted to one cell layer, which leads to a cell wall that is 10 times thicker for lenticels. They also displayed a different chemical composition because of unsuberization and a high lignin content in lenticels. Such advances in the knowledge of the structure and composition of cork cells contributes to a better understanding of the macroporosity of cork, down to the nanoscale.