Histochemical structure and tensile properties of birch cork cell walls

Tensile tests of birch cork were performed in the tangential direction. Birch cork in the wet state showed significantly higher extensibility than those in the dried state. The histochemical structure of birch cork was investigated by microscopic observation and spectroscopic analysis. Birch cork cell walls showed a two-layered structure and the inner material bordering cell wall. In transmission electron micrographs, osmium tetroxide stained the outer layer and inner material, whereas potassium permanganate stained the inner layer and inner material. After removal of suberin and lignin, only inner layer remained and Fourier-transformed infrared spectra showed the cellulose I pattern. Polarizing light micrographs indicated that molecular chains in the outer layer and inner material were oriented perpendicular to suberin lamination, whereas those in the inner layer showed longitudinal orientation. These results suggested that the outer layer and inner material mainly consist of suberin, whereas the inner layer and compound middle lamella consist of lignin, cellulose, and other polysaccharides. We hypothesized a hierarchical model of the birch cork cell wall. The lignified cell wall with helical arrangement of cellulose microfibrils is sandwiched between suberized outer layer and inner material. Cellulose microfibrils in the inner layer bear tensile loads. In the wet state, water and cellulose transfer tensile stress. In the dried state, this stress-transferal system functions poorly and fewer cells bear stress. Suberin in the outer layer and inner material may prevent absolute drying to maintain mechanical properties of the bark and to bear tensile stress caused by trunk diameter growth.