11th Class Biology Anatomy of Flowering Plants Secondary Growth

The increase in thickness or girth due to the activity of the cambium and the cork cambium is known as secondary growth.

(1) Secondary growth in stem : On the basis of the activities of cambium and cork-cambium, secondary growth in stem can be discussed under the following heads :

Activity of cambium : The vascular cambium in between xylem and phloem is called intrafascicular or fascicular cambium which is primary in origin. At the time of secondary growth the parenchymatous cells of medullary rays between the vascular bundles become meristematic and form strip of cambium called as interfascicular cambium which is secondary in origin. Both inter and intrafascicular cambium joins together and form cambium ring which is partly primary and partly secondary in origin. By anticlinal divisions the circumference of the cambium increase. By periclinal division cambium produces the secondary xylem and phloem tissues on innerside and outerside. The amount of sec. xylem produced is 8-10 times greater than sec. phloem. The cambium has two types of cells :

The fusiform initials : Which are elongated and form fibres, sieve cells, sieve tubes, tracheids.

Ray initials : Which produce parenchyma cells of the rays in wood and phloem. Certain cells of cambium form some narrow bands of living parenchyma cells passing through secondary xylem and secondary phloem and are called secondary medullary rays. These provide radial conduction of food from the phloem, and water and mineral salts from the xylem.

Annual rings : Activity of cambium is not uniform in those plants which grow in the regions where favourable climatic conditions (spring or rainy season) alternate regularly with unfavourable climatic conditions (cold water or dry hot summer). In temperate climates, cambium becomes more active in spring and forms greater number of vessels with wider cavities; while in winter it becomes less active and forms narrower and smaller vessels. The wood formed in the spring is known as spring wood and that formed in the dry summer or cold winter autumn wood or late wood. Both autumn and spring wood constitute a growth or annual ring. In one year only one growth ring is formed. Spring wood is light in colour while autumn wood is dark in colour.

Activity of cork cambium : Cork cambium or phellogen develops from outer layer of cortex. It produces secondary cortex or phelloderm on innerside and cork or phellem on outerside. The cells of phellem are dead, suberized and impervious to water. Cells of phelloderm are thin walled, living and store food. Phellem, phellogen and phelloderm are collectively called as periderm. Periderm is secondary protective tissue. Due to pressure of secondary xylem, epidermis ruptures and cortex is largely lost after two or three years of secondary growth.

Bark : All dead tissues lying outside the active cork-cambium are collectively known as bark. This includes ruptured epidermis, hypodermis and cork. When cork-cambium appears in the form of a complete ring, it is known as ring bark, e.g., Betula (Bhojpatra). If the cork cambium occurs as separate strips and the resulting bark appears in the form of scales, such a bark is known as scaly bark. e.g., Eucalyptus, Psidium guava. The outermost layer of bark is dead and called as rhytidome.

Lenticels : These are aerating pores formed in the cork through which gaseous exchange takes place. They are formed as a result of the action of phellogen. A lenticel appears as a scar or protrusion on the surface of the stem and consists of a radial row of thin-walled cells, known as complementary cells or filling tissue. They are found in old dicot stem, main function is gas exchange.

Cork : It consists of dead cells with thick walls heavily impregnated with suberin. These cells are compactly arranged in radial rows without intercellular spaces. Cork is impervious to water and prevents its loss from the plant surface. It also protects the inner tissues from the attack of fungi and insects. There is no differentiation of bark, sap wood and heart wood of Date palm.

Heart wood and sap wood : In old trees, secondary wood is differentiated into a centrally situated darker and harder wood called the heart wood or duramen which are physiologically inactive (almost dead)and an outer light-coloured zone called the sap wood or alburnum which are physiologically active. Dark colour of heart wood is due to the deposition of tannins, resins, gums, essential oils, etc. in the cell walls and cell cavities. The water conduction takes place through sap wood. During the conversion of sap wood into heartwood the most important change is development of tyloses in the heart wood. Tyloses are balloon like structures, develop from xylem parenchyma. These tyloses block the passage of xylem vessels so also called as tracheal plug. The heart wood is commercially used as wood. When the plant is made hollow, it will not die because the water conduction takes place through sap wood. The heart wood is well developed in Morus alba (Mulberry). The heart wood is absent in Populus and Salix plant. As a tree grows older thickness of heartwood increases and sap wood remains same.

(2) Secondary growth in dicot roots : Vascular bundles in dicot roots are radial, exarch and mostly triarch. Vascular cambium is formed secondarily from conjunctive parenchyma cells lying just below each phloem strand. Thus the number of cambium strips formed equals the number of phloem strands. The cells of pericycle lying outside the protoxylem also become meristematic to form part of strips of cambium. These cambial strips join the first formed cambium strips to form complete but wavy ring of vascular cambium. This cambium ring produces secondary xylem on inner side and secondary phloem on outer side. In roots, the growth rings are not distinct because there is no seasonal variation under the soil. From the outer layers of pericycle arises the phellogen which cuts phellem (cork) on the outer side and secondary cortex or phelloderm toward the inner side.