7.2.4 - The cork cambium

As the plant stem increases in girth during secondary growth, the outermost protective layer of the stem, the epidermis, is gradually shed and replaced by a periderm. Peridermal tissue is produced by the cork cambium (or phellogen) which, like the vascular cambium, comprises a meristematic layer that produces derivatives, both centrifugally and centripetally, and undergoes periclinal and anticlinal division forming radial files of cells. Derivatives from the phellogen differ from those of the cambium both structurally and functionally and the rate of division from this meristem is often much lower than that observed in the vascular cambium. The periderm is comprised of three different tissue types; 1) the phellogen or cork cambium, the meristematic tissue of the periderm, 2) phellum or cork, the outermost protective layer of tissue, and 3) phelloderm, the living parenchyma formed toward the inside of the phellogen. The phellogen is made up of a single layer or tier of cells that are radially flattened, rectangular to polygonal in shape, with thin primary walls, dense cytoplasm and, in some cases, chloroplasts. Phellum cells divide centrifugally from the phellogen and are highly specialised, creating an imperious barrier to gases and liquids on the plant stem. Phellum cells are usually polyhedral in shape, but can be radially compressed and/or elongated in several planes (radial, tangential and longitudinal) and are arranged compactly, lacking intercellular spaces. Phellum cells develop a layer of suberin, an unsaturated fatty acid that impregnates the cell wall during development and gives them their impervious properties. Maturation of these cells ultimately results in cell death. In some genera, e.g. Eucalyptus, phellum cells can also be lignified. Phelloderm cells are living parenchyma cells that divide centripetally from the phellogen and are similar in appearance to the adjacent cortical tissue, being only distinguishable by their radial alignment with cells in the phellogen.

The periderm is continuously replaced during growth and the site and tissue type from which it develops differ between species. Initiation of a periderm occurs either uniformly as a ring around the plant, or in scattered groups which link up later via lateral spreading.

Once the first periderm is ruptured, initiation of the next periderm usually occurs from either cortical tissue or phloem parenchyma whereas later periderms form exclusively from phloem parenchyma. This process is repeated continuously during the life of the plant. The formation of a new periderm displaces a large number of cells, which die and are shed from the stem, thus a tree continues to increase in diameter.