Organization of the vascular tissue

Vascular tissue is organized into discrete strands called vascular bundles, each containing xylem and phloem. In stems, the vascular tissue is organized into many discrete vascular bundles. In the roots, the vascular tissue is organized within a single central vascular cylinder. The anatomy of roots and stems is discussed in their respective sections below.

The xylem conducts water and minerals within the primary plant body, and the phloem conducts food. The xylem cells are arranged end to end to form a longitudinal continuum throughout the plant. The phloem cells form a similar continuum. Thus, water enters the xylem cells in the roots and travels to the leaves via the stems, and photosynthates (products of photosynthesis) enter the phloem cells in the leaves and are translocated to the roots via the stems. Storage parenchyma and fibres are generally present, and sclereids rarely are.

Primary xylem (Figure 6) consists of lignified tracheary elements (tracheids and vessel elements), which are dead at maturity (they have lost their protoplasts). Parenchyma cells also are interspersed throughout the tissue. Both tracheids and vessel elements are long hollow cells with tapered end walls. The end walls of adjacent tracheids contain paired small, rimmed, nonperforated pores, called bordered pits; water diffuses through a shared central membrane. The side walls have five patterns of thickening, which are believed to represent a developmental sequence from the initial xylem (protoxylem) to the final mature xylem (metaxylem): annular (a series of rings), helical (a long continuous spiral), reticular (a network), scalariform (a series of elongated bordered slits), and circular bordered pitting. Individual species may omit some of these patterns.

Vessel elements differ from tracheids in that the end walls are modified into perforation plates, an area or areas in which there is no shared wall material or membrane. Vessel elements join to form continuous vessels. The perforations are much larger than those of the bordered pits of tracheids and are of four types: scalariform (slitlike), foraminate (circular), reticulate (a network), or simple (single). The bordered pitting of the side walls of vessel members is either scalariform or circular (generally scalariform bordered pitting is associated with scalariform, foraminate, or reticulate perforation plates). Vessel elements are found in the late metaxylem (the final, or most developed, form of the primary xylem).

The most common type of perforation plates in the angiosperms are scalariform and simple; the other types are rare. The putatively primitive angiosperms are without vessels and evolved from a condition in which only tracheids were present to one in which a series of long vessel elements had scalariform lateral walls and highly inclined end walls with many scalariform perforations, to short vessel elements with circular bordered pits in lateral walls and simple perforation plates in horizontal end walls.

This series of specializations has increased the efficiency with which water moves through the vessels: from the more generalized method of water diffusion through pit membranes of narrow tracheids to mass movement of water through the perforated end walls of relatively narrow scalariform vessels and then to relatively wide simple vessels with large single perforated end walls. This simple form is a rather streamlined system that facilitates the maximum movement of water in terms of amount and speed with the minimum amount of resistance, allowing for greater efficiency and effective water transport.

The primary phloem (Figure 6) is composed of sieve elements and fibres. Parenchyma cells are interspersed throughout. Sieve elements are longitudinal cells that transport food. They are composed of sieve cells and sieve-tube members. Sieve-tube members have clusters of pores in the cell walls known as sieve areas, which have either small pores or large pores; the latter are known as sieve plates. Sieve plates are mostly located on the overlapping adjacent end walls. As sieve-tube members differentiate, they lose their nucleus, ribosomes, vacuoles, and dictyosomes (the equivalent of the Golgi apparatus in animals); they are not dead, however, and remain metabolically active. Each sieve-tube member has an associated specialized parenchyma cell called a companion cell. They are derived by mitosis from the same parent cell and remain connected with each other. Photosynthates are actively secreted into, and actively removed from, sieve-tube members by their companion cells. Other unspecialized parenchyma cells also are present in primary phloem and provide storage.

Finally, the primary vascular tissue system usually has fibres, particularly in herbaceous plants. The fibres occur in groups either around vascular bundles or as a cap over the phloem (phloem fibres).

The primary vascular system (Figure 7) serves three functions. First, the sieve tubes conduct photosynthates via companion cells from green stems and leaves to nongreen areas (usually roots, lateral meristems, and shoot apical meristems) to promote growth and development. Second, tracheary elements provide a water-conducting system and a support system as a result of their rigid lignified cell walls. Third, fibres provide additional support.

Secondary vascular system

In woody plants, a vascular system of secondary vascular tissue develops from a lateral meristem called the vascular cambium (Figure 8). The vascular cambium, which produces xylem and phloem cells, originates from procambium that has not completely differentiated during the formation of primary xylem and primary phloem. The cambium is thought to be a single row of cells arranged as a cylinder that produces new cells: externally the secondary phloem and internally the secondary xylem. Because it is not possible to distinguish the cambium from its immediate cellular derivatives, which also divide and contribute to the formation of secondary tissues, the cambium and its immediate derivatives are usually referred to as the cambial zone.

Unlike the apical meristems, which consist of a population of similar cells, the cambium consists of two different cell types; the fusiform initials and the ray initials. The fusiform initials are elongated tapering cells that give rise to all cells of the vertical system of the secondary phloem and xylem (secondary tracheary elements, fibres, and sieve cells and the associated companion cells). The ray initials are isodiametric cells—about equal in all dimensions—and they produce the vascular rays, which constitute the horizontal system of secondary tissues; this horizontal system acts in the translocation and storage of food and water.

The fusiform and ray initials of the cambium divide in a plane tangential to the surface of the stem, with the long axes of the fusiform and ray initials parallel to the long axis of the plant organ. The cambium generates xylem mother cells toward the inside and phloem mother cells toward the outside. These cells in turn continue to divide tangentially, producing new cells that add to the xylem and to the phloem. Divisions of the cambium cells and xylem and phloem mother cells do not result in the production of equal amounts of secondary xylem and secondary phloem; because the cambium produces more cells internally than externally, more secondary xylem is produced than secondary phloem. Because divisions in the fusiform and ray initials are primarily tangential, new cells are regularly arranged in well-defined radial rows, a characteristic pattern for secondary vascular tissues.

Divisions in the cambium not only produce secondary vascular tissues but also increase the circumference of the cambium. As new cells are continuously added to the inside of the cambium, the cambium increases laterally (in circumference) to keep pace with the circumferential growth of the stem. In some plants, this is accomplished simply by radial division of the fusiform and ray initials. In other plants, the mechanism for increasing cambial diameter or increasing the number of cambial cells is more complex. If cambial activity is extensive, the primary tissues lying outside the cambium, such as primary phloem, cortex, and epidermis, are crushed by the pressure of new secondary tissue growth or become torn and obliterated because they cannot accommodate the rapidly increasing diameter of the plant.

As growth proceeds, the cork cambium forms in living cells of the epidermis, cortex, or, in some plants, phloem and produces a secondary protective tissue, the periderm. The cork cambium is, like the vascular cambium, a lateral meristem that produces cells internally and externally by tangential divisions. Unlike the cambium, the cork cambium consists of one cell type.

Another type of meristem active in certain plants, especially grasses, is the intercalary meristem. These cells possess the ability to divide and produce new cells, as do apical and lateral meristems. They differ, however, in being situated between regions of mature tissue, such as at the base of grass leaves, which are themselves located on mature stem tissue. In many instances intercalary meristems function for only a short time and eventually completely differentiate into mature tissues. Intercalary meristems are usually located at positions on the stem where leaves have emerged (nodes) and are largely responsible for elongation in grass shoots and leaves. Intercalary meristems are the internode regions where cell division of the ground meristem persists for a longer time than in other areas of the internode. In rosette plants, intercalary meristems are lacking.

Secondary xylem is composed of tracheary elements, rays, fibres, and interspersed axial parenchyma cells. The tracheary elements consist of only tracheids, as in the few vessel-less angiosperms (e.g., Winteraceae), or of both tracheids and vessel elements, as in the vast majority of angiosperms. Axial parenchyma may surround the vessel elements (paratracheal) or be randomly dispersed among the vessel elements (apotracheal).

Tyloses are balloonlike outgrowths of parenchyma cells that bulge through the circular bordered pits of vessel members and block water movement. The presence of tyloses in white oaks makes their wood watertight, which is why it is preferred in casks and shipbuilding to red oak, which lacks tyloses and does not hold water. In trunks and branches that lean, there is eccentric growth of tension wood on the upper surface; tension wood is a type of reaction wood found in angiosperms that contains gelatinous fibres which shrink and pull.

Growth rings in the secondary xylem of temperate woody angiosperms are usually annual, but under environmental fluctuations, such as drought, more than one can form, or none at all. Growth rings result from the difference in density between the early wood (spring wood) and the late wood (summer wood); early wood is less dense because the cells are larger and their walls are thinner. Although the transition of early wood to late wood within a growth ring may be obscure, that demarcation between the adjacent late wood of one ring and the early wood of the next ring is clear. Diffuse-porous wood occurs when the size of the vessels (pores) in a growth ring are fairly uniform and evenly distributed (e.g., red maple, Acer rubrum; Sapindaceae). Ring-porous wood occurs when the pores of the early wood are distinctly larger than those of the late wood (e.g., black walnut, Juglans nigra; Juglandaceae).

Both xylem and phloem have limited longevity. The oldest phloem layers are the outermost—the dead bark of the stem surface. The yearly amounts of xylem visible as distinct rings in cross sections of stems are known as annual rings. The oldest xylem layers (i.e., the oldest annual rings) are in the dead central core, or heartwood, of the woody stem, which can often be recognized by its darker coloration. The lighter-coloured sapwood is living and functions as storage tissue and, especially in the outermost sapwood, as conducting tissue; the younger annual rings make up the sapwood. In some highly specialized tree species with large vessels (such as some oaks, ashes, and others), only the very outermost growth ring functions in water conduction.

Conducting tissues seldom run straight along a tree stem; usually they are arranged in a helical or spiral pattern, sometimes called the spiral grain of a tree. The angle of the spiral arrangement usually changes from year to year; the path of water up a tree stem may therefore be very complicated if more than one growth layer acts as a conducting tissue. Functionally, the effect of the variable spiral grain is to distribute water to all parts of the tree from any root.

The secondary phloem of angiosperms consists of sieve-tube members, companion cells, scattered parenchyma, ray parenchyma, and fibres. The fibres usually occur in clusters or as bands alternating with bands of sieve tubes and parenchyma cells. As the vascular cambium continues to produce more secondary xylem to the inside, the older (most exterior) portions of the secondary phloem are crushed, die, and are sloughed off as part of the bark. Successive cork cambiums (see below Dermal tissue), essentially lateral meristems from which the bark arises, originate in the parenchyma of the phloem and produce additional cork.

Uptake of water and mineral nutrients from the soil

Water uptake from the soil by root cells is passive, in that water may be pulled into the root by low xylem pressure and also follows osmotic gradients caused by the mineral nutrients, which are taken up actively (i.e., with the expenditure of metabolic energy) across root cell membranes. As the mineral nutrients—the ions (charged components) of inorganic salts—are taken up, they are largely incorporated into organic molecules. Thus, the solutes in xylem sap are mostly complex organic substances, sometimes of a specific nature; for example, nicotine synthesis takes place in the roots of tobacco plants, where nitrogen is incorporated into compounds that have moved to the roots through the phloem as sugars. If a tomato shoot is grafted onto a tobacco rootstock, nicotine-containing tomato leaves are formed. On the other hand, a tobacco shoot grafted onto a tomato rootstock results in a plant with nicotine-free tobacco leaves. Many other specific nitrogen-containing substances originate in the roots; in most plants, however, nitrogen is transported to the leaves from the roots in the form of compounds known as amino acids and amides.

The major chemical elements needed by a plant are carbon, hydrogen, oxygen, phosphorus, potassium, nitrogen, calcium, iron, and magnesium; in addition, many other elements are required in very small amounts. A lack of any element may result in deficiency diseases. A few elements taken up by plants are of no nutritive value and usually are eliminated or crystallized (e.g., silica), sometimes by deposition in special cells.

The plant is able to control to some extent the substances that enter. If equal amounts of sodium and potassium are available to roots of plants, and the amount of the two elements inside the plant is analyzed, less sodium is likely to be found than potassium. The structural basis for the control of uptake of substances into roots is the so-called Casparian strip, a conspicuously thickened wall area one cell layer deep surrounding primary roots; it prevents excess soil solution from being pulled directly into the central part of the root where the xylem is located. As a result, the soil solution has to pass through a cell barrier in which uptake can be metabolically controlled. After nutrients are inside living root cells and have been converted to appropriate compounds, the latter are released into the xylem and move to above-ground parts.