Structure and function

Vegetative structures

There are three levels of integrated organization in the vegetative plant body: organ, tissue system, and tissue. The organs of the plant—the roots, stems, and leaves—are composed of tissue systems (dermal tissue, ground tissue, and vascular tissue; see below Tissue systems). The tissues of each of these systems are composed of cells of one or more types (parenchyma, collenchyma, and sclerenchyma; see below Tissue systems: Ground tissue). Tissues composed of only one cell type and performing only one function are simple tissues, while those composed of more than one cell type and performing more than one function, such as support and conduction, are complex tissues. Xylem and phloem are examples of complex tissues.

The plant develops from a fertilized egg, called a zygote, which undergoes mitotic cell division to form an embryo—a simple multicellular structure of undifferentiated cells (i.e., those that have not developed into cells of a specific type)—and eventually a mature plant. The embryo consists of a bipolar axis that bears one or two cotyledons, or seed leaves; in most eudicots the cotyledons contain stored food in the form of proteins, lipids, and starch, or they are photosynthetic and produce these products, whereas in most monocots and some eudicots the endosperm stores the food and the cotyledons absorb the digested food. The embryos of eudicotyledons have two seed leaves, while those of monocotyledons have only one.

As the embryo continues to develop and new cells arise, the angiospermous plant develops specialized regions in which only cell division takes place and other areas in which nonreproductive (vegetative) activities, such as metabolism, respiration, and storage, occur. The areas of dividing cells, essentially permanently embryonic tissue, are called meristems, and their cells are termed initials. In the embryo they are found at either end of the bipolar axis and are called apical meristems. As the plant matures, apical meristems in the shoots produce new buds and leaves, and apical meristems in the roots are the points of active growth for roots (Figure 3). All growth produced by the apical meristems is primary growth and results in more primary tissues, which essentially extends the primary plant body.

After a cell in an apical meristem has divided mitotically, one of the two resulting daughter cells remains in the meristem as an initial cell, and the other cell is displaced into the plant body as a derivative cell. The displaced derivative cell may divide several times as it differentiates (changes in structure and physiology) from a meristemic cell into a mature cell, but only initial cells remain permanently in the apical meristem. However, although most permanently differentiated derivative cells are nondividing cells, and regions of division remain in the root and shoot apical meristems, there are regions of dividing derivative cells behind apical meristems that give rise to primary tissue systems and thus are also considered to be primary meristems.

Three concentric regions of primary meristematic tissues develop immediately behind the apical meristem (Figure 3). These primary meristems produce the different tissues of the plant body: the outermost protoderm differentiates into the epidermis, a tissue that protects the plant; the adjacent ground meristem differentiates into the central ground tissues (the pith and cortex); and the procambium differentiates into the vascular tissues (the xylem, phloem, and vascular cambium). The xylem and phloem are conducting and supporting vascular tissues, and the vascular cambium is a lateral meristem that gives rise to the secondary vascular tissues, which constitute the secondary plant body.

Lateral meristems, called cambia, run the length of the stems and roots of vascular plants and produce secondary tissues, which develop after a plant organ—or part of a plant organ—has ceased to elongate. Secondary growth is essentially an increase in girth. The vascular cambium produces secondary xylem and secondary phloem, and the cork cambium (phellogen) produces cork cells, from which the outer bark develops. Figure 4 summarizes the patterns of primary and secondary growth from root and shoot apical meristems. (For a complete discussion of primary and secondary plant growth, see below Tissue systems: Vascular tissue.)

Tissue systems

As mentioned above, three areas of meristematic tissue are derived directly from the apical meristem: the ground meristem, procambium, and protoderm. These meristematic tissues differentiate into the three primary tissues that constitute the primary plant body: ground tissue (pith and cortex), vascular tissue (xylem, phloem, and eventually the lateral, or secondary, meristem called the vascular cambium), and dermal tissue (epidermis), respectively.