Delphinidin

A novelty song of more than 50 years ago listed food items the writer said he disliked. Each verse ended with the line, “I like bananas because they have no bones!” Indeed, bananas and all plants differ from larger animals in having no bones or similar internal skeletal structures. Yet large trees support branches and leaves weighing many tons. They can do this because most plant cells have either rigid walls that provide the support afforded to animals by bones or semi-rigid walls that provide flexibility. At the same time, the walls protect delicate cell contents within. When millions of these cells function together as a tissue, their collective strength is enormous. The redwoods and Tasmanian Eucalyptus trees, which are the largest trees alive today, exceed the mass and volume of the largest land animals, the elephants, by more than a hundred times. The wood of one giant redwood tree could support the combined weight of a thousand elephants.

The first cell structure discovered by Robert Hooke in 1665 was the cell wall, and among plant cell structures observed with a microscope, the cell wall is the most obvious because it defines the shape of the cell. Many of the prepared specimens observed with a microscope in plant biology are merely stained remnants of once-living cells. But the vast diversity of cell walls within and among species tells a story about the structure and function of each cell. For instance, epidermal cells, which form a thin layer on the surfaces of all plant organs, often have unusual shapes and sizes. Some such cells form hairs that may secrete substances that discourage animals from grazing on the plants producing them. Thin-walled cells found beneath the epidermis of leaves are specialized for their function of
photosynthesis; and thick-walled cells of wood help to transport water without collapsing.

The main structural component of cell walls is cellulose, which is composed of 100 to 15,000 glucose monomers in long chains, and is the most abundant polymer on earth. As a primary food source for grazing animals and at least indirectly for nearly all other living organisms, it could be said that most life on earth relies directly or indirectly on the cell wall. Humans also depend on cell walls because they provide clothing, shelter, furniture, paper, and fuel.

In addition to cellulose, cell walls typically contain a matrix of hemicellulose (a gluelike substance that holds cellulose fibrils together), pectin (the organic material that gives stiffness to fruit jellies), and glycoproteins (proteins that have sugars associated with their molecules).

A middle lamella, which consists of a layer of pectin, is first produced when new cell walls are formed. This middle lamella is normally shared by two adjacent cells and is so thin that it may not be visible with an ordinary light microscope unless it is specially stained. A flexible primary wall, consisting of a fine network of cellulose, hemicellulose, pectin, and glycoproteins, is laid down on either side of the middle lamella. Reorganization, synthesis of new molecules, and insertion of new wall polymers lead to rearrangement of the cell wall during growth. Secondary walls, which are produced inside the primary walls, are derived from primary walls by thickening and inclusion of lignin, a complex polymer.

Secondary cell walls of plants generally contain more cellulose (40% to 80%) than primary walls. As the cell ages, wall thickness can vary, occupying as little as 5% to more than 95% of the volume of the cells. During secondary wall formation, cellulose microfibrils become embedded in lignin, much like steel rods are embedded in concrete to form prestressed concrete.