How wood & bark develop How stems & roots become thicker and stronger Secondary growth • Secondary growth> is growth in thickness in gymnosperms & angiosperms • Occurs in roots and stems of woody plants. • Requires first the establishment of secondary meristems (also called lateral meristems) • Results in from cell divisions in the vascular & cork cambium • Vascular cambium produces secondary xylem & sec. phloem • Cork cambium produces the layers of cork • Sec. xylem & sec. phloem function just like the primary xylem & phloem, by moving water, minerals & food • Secondary xylem and secondary phloem provide physical continuity between primary xylem and phloem of roots and stems • As new vascular tissue is formed, the root, stem or branch increases in diameter • The tissue located towards the outside of the vascular cambium compose the bark of a tree • Main products are wood (sec xylem) and bark (sec phloem) - the periderm (replaces epidermis). Secondary growth in Roots Secondary growth in roots The secondary meristems involved are: 1. Vascular cambium divides to produce secondary xylem to the inside and secondary phloem to the outside 2. Cork cambium or phellogen divides to produce phelloderm to the inside and cork to the outside Sec. growth in roots cont… • The vascular cambium (one cell layer thick) differentiates from the procambium (between the primary xylem and primary phloem) and from the pericycle at the ridges of the primary xylem where the protoxylem is in contact with the pericycle
• This results in an smoothly continuous vascular
cambium around the primary xylem Sec. growth in roots cont… • Formation of sec. xylem & sec. phloem results in increased thickness of the root
•The cortex and epidermis become broken, and
shed off, since their cells cannot divide, ultimately the root is covered by the cork on the outside
•As these tissues break, the pericycle cells are
induced to divide and produce a single layer of meristematic cells, the cork cambium or phellogen Formation of the periderm •The phellogen divides to produce the other type of secondary tissue, the periderm.
•Periderm is formed from cork cambium/phellogen
•Periderm replaces epidermis in sec. growth
•The phellogen divides to produce cork or phellem
to the outside and phelloderm to the inside. Formation of periderm cont… • 3 layered 1) Phellem /cork (outer layer - several layers thick) 2) Phellogen /cork cambium (middle layer- one cell layer thick) 3) Phelloderm (inner layer - about 3 layers thick)
• The phellogen (middle layer) is initiated
immediately after the formation of the vascular cambium Secondary growth in Stems Secondary growth in stems • The involved meristems are the vascular cambium and phellogen/cork cambium.
• Procambial cells between primary xylem and
primary phloem form bundle part of the vascular cambium – fascicular cambium. At this stage the vascular cambium is discontinuous.
• Some parenchyma cells of the pith ray that are in
line with the fascicular cambium differentiates to form another part of VC- interfascicular cambium Sec. growth in stems cont… • The vascular cambium consists of fusiform initials (elongated cells) and ray initials (isodiametric cells)
• The fusiform initials form the axial system
(conductive cells) and the ray initials form the radial system, parenchyma rays that function in storage and lateral transport • Xylem rays> consist of parenchyma cells & tracheids, • Phloem rays> consist of parenchyma cells Sec. growth in stems cont… • The vascular cambium divide by periclinal divisions to produce secondary phloem to the outside and secondary xylem to the inside • The secondary xylem is wood; wood formed in one year appear as one ring – the annual ring • Each annual ring consists of early growth (spring wood) and late growth (autumn wood) • Spring/early wood occurs to the inside, has tracheary elements of larger diameter with thinner walls • Autumn/late wood consists of tracheary elements of smaller diameter with thicker walls Sec. growth in stems cont.. • Wood is characterised by type of cells that compose it • Ring porous wood: wood in which tracheary elements with large diameter occurs only in the spring wood • Diffuse porous wood: wood in which tracheary elements with large diameter are distributed evenly in the spring and autumn wood
• Heartwood: wood that occur at the centre of the
stem, is dark in colour, it is nonfunctional (blocked by tyloses) • Sapwood: wood that occur towards the outside, it Heartwood Sapwood
Colour Dark Lighter
Alive/dead Dead (non-functional) Functional
Function Supports the tree Supports the tree
Occurrence In the central portion of an old In the outer portion of an old
stem stem
Durability Durable, furniture Not durable
Weight Heavier Lighter
Sec. growth in stems cont.. • Cork has three layers – phellem, phellogen and phelloderm • Cork is often confused with bark…bark referes to all tissues outside the VC-mostly the non-wood of the stem • Cork cambium/phellogen forms from parenchyma cells on the outer layer of the cortex; a single layer of cells. • As the wood increases in diameter during growth, the bark cannot keep up with the rate of growth. Sec. growth in stems cont.. • Since cork is part of bark, when bark brakes down it exposes the plant to microorganisms.
• To maintain protection, new phellogen then
differentiates from parenchyma cells of the secondary phloem
• In some plants, new periderms are formed
internally from secondary phloem, whilst the older, outer periderms still exist to form a rhytidome – a mixture of dead and living layers Fig. 1 • The laterally positioned meristems divide to produce one type of secondary tissue to the inside and another type of tissue to the outside. (Fig. 1)