Secondary Growth

• The growth of the roots and stems in length with the help of apical meristem is called the primary growth.

• Apart from primary growth most dicotyledonous plants exhibit an increase in width. This increase is called the secondary growth. 

The tissues involved in secondary growth are the two lateral meristems:

· Vascular cambium

· Cork cambium

Vascular Cambium:

• The meristematic layer that is accountable for severing off vascular tissues – xylem and pholem – is called vascular cambium.

• In the young stem it is present in patches as a single layer between the xylem and phloem and forms a complete ring, in the late stages.

Formation of Cambial Ring:

• In dicot stems, the cells of cambium present between primary xylem and primary phloem is the intrafascicular cambium. The cells of medullary rays, adjoining these intrafascicular cambium become meristematic and form the interfascicular cambium. Thus, a continuous ring of cambium is formed.

Activity of the Cambial Ring:

• The cambial ring becomes active and starts to remove new cells, both on the inner and outer sides. The cells cut off towards pith, mature into secondary xylem and the cells cut off towards periphery mature into secondary phloem.

• The cambium is generally more active on the inner side than on the outer. As a result, the amount of secondary xylem produced is more than secondary phloem and soon forms a compact mass.

• The primary and secondary phloems get gradually crushed due to the continued formation and accumulation of secondary xylem. The primary xylem however remains more or less intact, in or around the centre. At some places, the cambium forms a narrow band of parenchyma, which passes through the secondary xylem and the secondary phloem in the radial directions. These are the secondary medullary rays. 

Secondary growth in a dicot stem (diagrammatic) – stages in transverse views

Spring Wood and Autumn Wood:

• Many physiological and environmental factors, controls the activity of cambium. In moderate regions, the climatic conditions are not uniform for the entire year. 

• In the spring season, cambium is very active and produces a large number of xylary elements having vessels with wider cavities. The wood formed during this season is called spring wood or early wood.

• In winter, the cambium is less active and forms fewer xylary elements that have narrow vessels, and this wood is called autumn wood or late wood. 

• The spring wood is lighter in colour and has a lower density whereas the autumn wood is darker and has a higher density. 

• The two kinds of woods that look as alternate concentric rings, constitute an annual ring. An estimate of the age of the tree can be done from the Annual rings seen in a cut stem.

Heartwood and Sapwood:

• In old trees, the greater part of secondary xylem is dark brown due to deposition of organic compounds like tannins, resins, oils, gums, aromatic substances and essential oils in the central or innermost layers of the stem. These substances make it hard, durable and resistant to the attacks of microorganisms and insects. This region comprises dead elements with highly lignified walls is called heartwood.

• The heartwood gives mechanical support to the stem, does not conduct water. The peripheral region of the secondary xylem, is lighter in colour and is known as the sapwood. It conducts water and minerals from root to leaf.

Cork Cambium:

• The activity of vascular cambium, the outer cortical and epidermis layers get ruined which makes the stem continues to increase in girth and need to be replaced to provide new protective cell layers. Immediately or later another meristematic tissue called cork cambium or phellogen develops, usually in the cortex region. 

• Phellogen is a couple of layers thick, it is made of narrow, thin-walled, nearly rectangular cells and it cuts off cells on both sides. The outer cells differentiate into cork or phellem while the inner cells differentiate into secondary cortex or phelloderm

• The cork is water-resistant due to suberin deposition in the cell wall. The cells of secondary cortex are parenchymatous. 

• Phellogen, phellem, and phelloderm are collectively known as periderm. 

• Due to activity of the cork cambium, pressure builds up on the remaining layers peripheral to phellogen and ultimately these layers die and slough off.

• All tissues exterior to the vascular cambium, including secondary phloem refers to bark and it denotes to a number of tissue types, viz., periderm and secondary phloem. Bark formed early in the season is early or soft bark. Late in the end of the season, late or hard bark is formed. 

Bark

• At certain regions, the phellogen cuts off closely arranged parenchymatous cells on the outer side in its place of cork cells. These parenchymatous cells soon rupture the epidermis, forming a lens shaped openings called lenticels. Lenticels permit the exchange of gases between the outer atmosphere and the internal tissue of the stem. These occur in most woody trees. 

Lenticel

Secondary Growth in Roots:

• In the dicot root, the vascular cambium is completely secondary in origin. It originates from the tissue located just below the phloem bundles, a portion of pericycle tissue, above the protoxylem forming a complete and continuous wavy ring, which later becomes circular.

Different stages of the secondary growth in a typical dicot root

• Secondary growth also occurs in stems and roots of gymnosperms. However in monocotyledons secondary growth does not occur.