Secondary growth in dicot stems occurs through the formation of a cambium ring by the vascular cambium and ray initials. This cambium ring produces secondary phloem and secondary xylem tissues on its outer and inner sides respectively through periclinal cell divisions. The secondary xylem accumulates to form wood over time, and along with seasonal variations in its formation, produces growth rings that can indicate a tree's age. In older stems, the secondary xylem differentiates into non-living heartwood at the center and living sapwood nearer the phloem.
Secondary growth in dicot stems occurs through the formation of a cambium ring by the vascular cambium and ray initials. This cambium ring produces secondary phloem and secondary xylem tissues on its outer and inner sides respectively through periclinal cell divisions. The secondary xylem accumulates to form wood over time, and along with seasonal variations in its formation, produces growth rings that can indicate a tree's age. In older stems, the secondary xylem differentiates into non-living heartwood at the center and living sapwood nearer the phloem.
Secondary growth in dicot stems occurs through the formation of a cambium ring by the vascular cambium and ray initials. This cambium ring produces secondary phloem and secondary xylem tissues on its outer and inner sides respectively through periclinal cell divisions. The secondary xylem accumulates to form wood over time, and along with seasonal variations in its formation, produces growth rings that can indicate a tree's age. In older stems, the secondary xylem differentiates into non-living heartwood at the center and living sapwood nearer the phloem.
The growth in length of main axis by the activity of apical meristem is called primary growth. Increase in thickness or girth of the axis due to the formation of secondary tissue is called secondary growth. The secondary tissue is formed by the activity of cambium and cork- cambium. The cambium forms secondary tissues in the stelar region and cork-cambium form secondary tissues into cortical regions. Secondary growth occurs only in dicot stem and root. It is usually absent in monocot root and stem.
QUESTIONS: WHAT IS THE STELAR REGION? WHAT IS THE CORTICAL REGION?
In dicot stem, secondary growth takes place through the following steps: a) Formation Of Cambium Ring Formation of the cambium ring is the first step of secondary growth. The cambium of vascular bundles becomes meristematic. At the same time some of the medullary ray cells lying at the level of cambium also become meristematic and form a strip of interfasciular cambium together with intrafasciular cambium form a complete circular ring, which is called cambium ring. The cambium ring forms the secondary tissues in the stelar region.
The vascular cambium consists of two types of cells, the fusiform initials and the ray initials. The fusiform initials are vertically oriented and divide to form the elements of xylem and phloem. The cells of ray initials are smaller and isidiametric which give rise to vascular rays of parenchymatous cells. b) Formation Of Secondary Vascular Tissues: The cambium ring cuts off new cells, both on outer and inner sides. The new cells formed on the outer side gradually modify into the elements of secondary phloem. The cells formed on the inner side gradually modify into secondary xylem.
Secondary Phloem: They consist of sieve tubes, companion cells, phloem parenchyma and phloem fibres. The primary phloem present on the outside gets crushed and is represented by small patches.
Secondary Xylem: Secondary xylem consists of vessels, tracheids, wood fibres and wood parenchyma. The vessels or trachea are most abundant in secondary xylem and are usually shorter than that of the primary xylem. The cambium ring forms more tissue on the inner side than on the outer side. As a result, secondary xylem forms the main bulk of the plant body and is generally called the wood. Its width increases with age. The primary xylem persists as conical projections towards the pith.
c) Vascular rays: Ray initials of the cambium ring form some narrow bands of parenchymatous cells. These cells extend radially from the pith to the phloem. These are called secondary medullary rays or vascular rays. The rays present in xylem are xylem rays and the rays present in phloem are called phloem rays. d) Annual rings: The activity of vascular cambium is greatly affected by the variations in the climate. It is more pronounced in temperate regions. The cambium stops dividing in winter. In the spring season or early summer, the cambium becomes more active and produces a large number of vessels with wider lumen. These are called spring wood or early wood. During the autumn or winter season, the cambium becomes less active and produces vessels with narrow lumens. Tracheids and wood fibres are formed in large numbers. These woods are called autumn wood or late wood. Hence, the annual rings are formed year after year. In the oldest part of the tree, annual rings can be used in determining the age of a tree.
In tropical regions, the climate is more or less uniform. Therefore, the annual rings are not well developed and do not correlate with the age of tree.
e) Sapwood and Heartwood:
In older stems, the woody trunk is differentiated into two regions. The outer light coloured region is called sap wood or alburnum and central dark colored region is called heart wood or duramen. The cells of sapwood are living and functional. They take part in conduction of water and storage of food. The heartwood consists of dead cells. During the growth process, the rings of sap wood gradually process, the rings or sap wood gradually convert into heartwood. The living cells of sap wood lose their protoplast and water content. The lumen of the xylem vessels get blocked by the ingrowth of the parenchyma cells. The adjacent parenchyma tissue enters through the pits of vessels and gradually enlarges to form a balloon like structure, which is called tyloses. The heartwood is stronger and more durable than sapwood. The heartwood becomes resistant to the attacks of bacteria and fungi due to the presence of antiseptic oils.
Secondary Growth in Dicot Roots
At completion of primary growth (i.e. growth in length), the secondary growth (i.e. growth in thickness) is initiated in roots due to the activity of the vascular cambium. Secondary growth includes the production of secondary phloem and secondary xylem by this lateral meristem. Roots with secondary growth often become woody due to the accumulation of secondary xylem. If we examine older thick roots of trees, we observe annual growth rings in the wood (secondary xylem) resembling those of stems.
The vascular cambium appears first on the inner edges of the phloem strands. While these cambial cells form some secondary elements, the pericyclic cells outside the protoxylem poles also divide. The inner derivatives of these divisions complete the cylinder of cambium by joining the strips located on the inner faces of the phloem strands. The vascular cambium loses its wavy circular outline in transverse sections because on the inner boundary of the phloem, the secondary xylem is deposited earlier than outside the protoxylem. The secondary vascular tissues assume the form of a continuous cylinder and complete enclose the primary xylem. The sieve elements of the primary phloem are crushed, and some of the remaining cells differentiate into fibers. The cambium which arises in the pericycle outside the xylem poles forms wide vascular rays. In a few species, the phloem remains internal to the xylem.
The initiation of phellogen and periderm follows the initiation of vascular cambium and secondary vascular tissues. Phellogen (cork cambium) originates from the pericycle cells located opposite primary phloem. The primary tissues located outside the pericycle (rhizodermis, exodermis, cortex and endodermis) become isolated from the secondary body of the root by periderm formation and die.
The secondary phloem in roots consists of axial elements such as sieve tubes with companion cells, phloem parenchyma cells and phloem fibers, and radial elements, i.e. ray cells. The axial elements of secondary xylem which are deposited by fusiform initials include wood parenchyma, fibers and vessels of various widths with bordered pits arranged in reticulate or scalariform patterns in their lateral walls. The rays are usually wide in herbaceous dicots and as narrow as in stems in woody life-forms. The root secondary xylem typically contains more parenchyma elements than the stem wood.
Periclinal divisions in the pericycle that are not involved in the formation of the vascular cambium occur not only outside the xylem poles, but spread around the circumference of the root. Such divisions are preparatory for the formation of the periderm. A phellogen arises among the outer cells of the proliferated pericycle which forms cork tissue to the outside and phelloderm towards the inside. The phelloderm is difficult to distinguish from the parenchyma derived from the pericycle.
Some dicotyledonous roots may retain their cortex for a period of time during secondary development. Such roots may develop an exodermis or a superficial periderm, and the cortex later becomes lost.
In some cases, the endodermis of the parent root takes part with the pericycle in forming branch roots. Sometimes it forms several cell layers by undergoing both anticlinal and periclinal divisions. In a few cases (e.g. Cucurbitaceae and certain water plants), the innermost cortical cell layers are also involved in the formation of the lateral root by contributing cells to the lateral primordium.
Adventitious roots may be formed in young organs or in older tissues that have not quite lost their meristematic properties. Most adventitious roots arise endogenously, sometimes from primordia laid down previously and remaining dormant until stimulated to growth. Usually the adventitious roots arise from the pericycle, but may also arise from the cambial zone.
Roots in both primary and secondary states of growth may undergo modifications in their structure due to the acquisition of special functions. In this sense, storage, aerial, mycorrhizal and contractile roots as well as roots modified by the interaction with parasitic plants are specially considered in this unit of study.
Many roots may form a symbiotic association with fungi termed mycorrhizae. In ectomycorrhizae, the fungal hyphae grow as a mantle around root endings and between the cells of the rhizodermis and cortex, while in endomycorrhizae the hyphae invade the root cells and the mantle is poorly developed.
Characteristics of Meristematic Tissues
The cells of the meristematic tissues are quite distinct in their cytological and physiological characteristics from other cells. The following are the characteristics of the meristematic cells.
1. Cells have the power of active division. 2. They are compactly arranged in tissue and there is absolutely no intercellular space. 3. The cell wall is thin and primary in nature, containing only cellulose. It is uniformly thick. There is no secondary thickening. 4. Cells possess dense protoplasm with a prominent large nucleus compared to other cells of equal volume. 5. Vacuoles are small or absent totally. 6. Cells do not possess ergastic sub-stances. Ergastic substances are non-protoplasm materials found in cells. 7. Mitochondria and endoplasmic reticulum are very little differen-tiated. 8. Plastids, when present, are in protoplastid stage. 9. Cells contain relatively more number of ribosomes. 10. Cells are metabolically very active.
When cells of the meristem divide, the daughter cells get differentiated into mature types while the others re-main meristematic. For this reason, the meristems perpetuate and become con-tinuous source of cell formation.
Function of the Vascular Cambium
Vascular Cambium produces secondary growth, giving rise to secondary xylem and secondary phloem.
The vascular cambium is a lateral meristem in the vascular tissue of plants. It is a cylinder of unspecialized meristematic cells that divide to give rise to cells that further divide, differentiate, and specialize to form the secondary vascular tissues.
The vascular cambium usually consists of two types of cells: Fusiform initials (tall cells, axially oriented); Ray initials (almost isodiametric cells, smaller and round to angular in shape).
Secondary xylem is the xylem that is formed during secondary growth from vascular cambium.
Phloem is the living tissue that carries organic nutrients (known as photosynthate), in particular, sucrose, a sugar, to all parts of the plant where needed.
The phloem is concerned mainly with the transport of soluble organic material made during photosynthesis.
Phloem
A vascular tissue in land plants primarily responsible for the distribution of sugars and nutrients manufactured in the shoot.
Xylem
A vascular tissue in land plants primarily responsible for the distribution of water and minerals taken up by the roots; also the primary component of wood.
Anomalous Secondary Growth
Anomalous secondary growth does not follow the pattern of a single vascular cambium producing xylem to the inside and phloem to the outside. Some dicots have anomalous secondary growth, such as Bougainvillea, where a series of cambia arise outside the oldest phloem. Most monocots either have no secondary growth or else anomalous secondary growth of some type. For example, palm trees increase their trunk diameter due to division and enlargement of parenchyma cells, which is termed "diffuse secondary growth." In some other monocot stems with anomalous secondary growth, a cambium forms, but it produces vascular bundles and parenchyma internally and only parenchyma externally. Some monocot stems increase in diameter due to the activity of a primary thickening meristem, which is derived from the apical meristem. Formation of Cork Cambium Cork originates from a layer of cambium (=phellogen) that itself is formed as a secondary meristem from a layer of collenchyma or parenchyma immediately beneath the epidermis. In contrast to sclerenchyma cells, collenchyma cells are alive and they have retained the potency to de-differentiate. Cork cambium cells only divide periclinally so that the typical rows of daughter cells arise: cork cells (=phellem) are mainly generated toward the outside. In a lesser extent also cork parenchyma (=phelloderm) is made toward the inside. Cork cambium, cork cells and cork parenchyma together are also named periderm. Mature cork cells are dead; their cell walls contain suberine, a fatty substance that repels water. The layer of cork provides protection against desiccation, but it also isolates tissues in the inner parts of the stem or trunk so thoroughly that exchange of gas with the outer world is impeded. 'Breathing' is yet achieved by so-called lenticels. The cork cambium starts to generate numerous parenchyma cells toward the surface, in most cases at the level of a stoma. These thin-walled parenchyma cells, which eventually degenerate, force an interruption of the sealing created by the cork layer and focus an opening for gas-exchange. Lenticel formation Lenticels originate beneath stomata, simultaneous with the initiation of the first layer or periderm or just before the initiation of periderm, during first growing season. As the lenticels formation starts, the parenchyma cells found near substomatal cavity lose their chlorophyll content and irregularly divide in different plants giving rise to a mass of colourless, rounded, thin walled, loosely arranged cells called Complementary cells. Sometimes complementary cells produced by phellogen towards outside instead of producing cork cells. As the complementary cells increase in number, pressure is exerted against the epidermis and it ruptures. Outer most cells gradually become dead and may be replaced. Beneath the outer layer some mass of closely packed cells alternate to loosely arranged cells are formed called as Closing layer. From inner side continuous production of complementary cells cause rupture of closing layer at intervals. Function:- i) Lenticels contain profuse inter cellular spaces for which it perform the function of exchange of gases between the atmosphere and internal tissue of the stem. ii) Lenticels like Stomata help in transpiration. Called Lenticelar transpiration. iii) Lenticels are active during night when stomatal transpiration stops. Periderm The periderm is the secondary protective (dermal) tissue that replaces the epidermis during growth in thickness of stems and roots of gymnosperms and dicotyledons (i.e., secondary growth). Unlike the epidermis, the periderm is a multilayered tissue system, the bulk of which usually constitutes the cork, or phellem. There are, however, some exceptions to this inasmuch as some other structures (e.g. potato skin and apple peel) are also periderm. The lateral meristem, (cork cambium or phellogen), is one cell layer thick and encircles the stem. It produces periderm centrifugally. The layer of cork cells formed is impermeable for water and gases, but is interrupted at certain points by lenticels which function to some extent similar to stomata in the epidermis, and permit gas diffusion. In some cases parenchyma cells are produced centripetally (i.e. to the inside of the stem or root) by the phellogen as a part of the periderm. These persistent living cells are called phelloderm and structurally appear similar to cells of the cortex. The number of layers of cork and phelloderm varies greatly among different species; some plants produce no phelloderm. The most important function of the periderm is to reduce the loss of water and solutes from interior tissues and to protect a plant from unfavorable environmental conditions.