Stem Growth

Left illustration shows a cross section of a woody stem undergoing primary growth. At the core of the stem is pith. Toward the outside are egg-shaped vascular bundles. Xylem is located toward the inside of the vascular bundle, and phloem is in the middle. Sclerenchyma cap the outside of the bundle. Right illustration shows a cross section of a woody stem undergoing secondary growth. As in primary growth, the core of the stem is pith. Outside the pith is a ring of secondary xylem. Rounded bundles of primary xylem tissue project from this ring into the pith. Outside the secondary xylem is a ring of secondary phloem tissue. The vascular cambium separates the xylem from the phloem. Outside the secondary phloem is the cortex layer. Bundles of primary phloem project outward from the secondary phloem into the cortex. A cork ring surrounds the cortex. The cork is separated from the cortex by a thin cork cambium. The bark of the tree extends from the vascular cambium to the epidermis.

Figure 1. In woody plants, primary growth is followed by secondary growth, which allows the plant stem to increase in thickness or girth. Secondary vascular tissue is added as the plant grows, as well as a cork layer. The bark of a tree extends from the vascular cambium to the epidermis.

Growth in plants occurs as the stems and roots lengthen. Some plants, especially those that are woody, also increase in thickness during their life span. The increase in length of the shoot and the root is referred to as primary growth, and is the result of cell division in the shoot apical meristem. Secondary growth is characterized by an increase in thickness or girth of the plant, and is caused by cell division in the lateral meristem. Figure 1 shows the areas of primary and secondary growth in a plant. Herbaceous plants mostly undergo primary growth, with hardly any secondary growth or increase in thickness. Secondary growth or “wood” is noticeable in woody plants; it occurs in some dicots, but occurs very rarely in monocots.

Some plant parts, such as stems and roots, continue to grow throughout a plant’s life: a phenomenon called indeterminate growth. Other plant parts, such as leaves and flowers, exhibit determinate growth, which ceases when a plant part reaches a particular size.

Primary Growth

Most primary growth occurs at the apices, or tips, of stems and roots. Primary growth is a result of rapidly dividing cells in the apical meristems at the shoot tip and root tip. Subsequent cell elongation also contributes to primary growth. The growth of shoots and roots during primary growth enables plants to continuously seek water (roots) or sunlight (shoots).

The influence of the apical bud on overall plant growth is known as apical dominance, which diminishes the growth of axillary buds that form along the sides of branches and stems. Most coniferous trees exhibit strong apical dominance, thus producing the typical conical Christmas tree shape. If the apical bud is removed, then the axillary buds will start forming lateral branches. Gardeners make use of this fact when they prune plants by cutting off the tops of branches, thus encouraging the axillary buds to grow out, giving the plant a bushy shape.

Watch this BBC Nature video showing how time-lapse photography captures plant growth at high speed.

Secondary Growth

The increase in stem thickness that results from secondary growth is due to the activity of the lateral meristems, which are lacking in herbaceous plants. Lateral meristems include the vascular cambium and, in woody plants, the cork cambium (see Figure 1).

Photo shows rough, white ovals embedded in a smooth, reddish brown woody tree trunk. Where the ovals are, it appears as if the bark has been scraped away.

Figure 2. Lenticels on the bark of this cherry tree enable the woody stem to exchange gases with the surrounding atmosphere. (credit: Roger Griffith)

The vascular cambium is located just outside the primary xylem and to the interior of the primary phloem. The cells of the vascular cambium divide and form secondary xylem (tracheids and vessel elements) to the inside, and secondary phloem (sieve elements and companion cells) to the outside. The thickening of the stem that occurs in secondary growth is due to the formation of secondary phloem and secondary xylem by the vascular cambium, plus the action of cork cambium, which forms the tough outermost layer of the stem. The cells of the secondary xylem contain lignin, which provides hardiness and strength.

In woody plants, cork cambium is the outermost lateral meristem. It produces cork cells (bark) containing a waxy substance known as suberin that can repel water. The bark protects the plant against physical damage and helps reduce water loss. The cork cambium also produces a layer of cells known as phelloderm, which grows inward from the cambium. The cork cambium, cork cells, and phelloderm are collectively termed the periderm. The periderm substitutes for the epidermis in mature plants. In some plants, the periderm has many openings, known as lenticels, which allow the interior cells to exchange gases with the outside atmosphere (Figure 2). This supplies oxygen to the living and metabolically active cells of the cortex, xylem and phloem.

Annual Rings

Photo shows a cross section of a large tree trunk with many rings projecting outward from the center.

Figure 3. The rate of wood growth increases in summer and decreases in winter, producing a characteristic ring for each year of growth. Seasonal changes in weather patterns can also affect the growth rate—note how the rings vary in thickness. (credit: Adrian Pingstone)

The activity of the vascular cambium gives rise to annual growth rings. During the spring growing season, cells of the secondary xylem have a large internal diameter and their primary cell walls are not extensively thickened. This is known as early wood, or spring wood. During the fall season, the secondary xylem develops thickened cell walls, forming late wood, or autumn wood, which is denser than early wood. This alternation of early and late wood is due largely to a seasonal decrease in the number of vessel elements and a seasonal increase in the number of tracheids. It results in the formation of an annual ring, which can be seen as a circular ring in the cross section of the stem (Figure 3). An examination of the number of annual rings and their nature (such as their size and cell wall thickness) can reveal the age of the tree and the prevailing climatic conditions during each season.