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10.1 10.2 10.3
Secpndary Growth in Typical Dicotyledonous Stem. Vascular Cambium-General Development and Structure
10.3.1 Structure of Vascular Cambium 10.3.2 Types of Cambium 10.3.3 Structure of Protoplast and Cell Growth 10.3.4 Ray Ipitiation.
10.4.1 Formation of Annual Rings.
10.5.1 Basic Structure of Secondary Xylem 10.5.2 Wood Parenchyma 10.5.3 Heart Wood and Sapwood 10.5.4 Economic importance of Wood and its Characteristics.
10.6 10.7 10.8
10.6.! Economic Value of Secondary Phloem.
Secondary Growth in Monocot Stem Periderm
10.8.1 Structure 10.8.2 Phellem 10.8.3 Phelloderm 10.8.4 Origin and Development of Peridenn 10.8.5 Commercial Cork
Distribution of Lenticels
10.9.1 Development and Structure of Lenticels
10.10 Cqmbial Variants
10.10.1 In Stems 10.10.2 In Roots.
10.11 Summary 10.12 Terminal Questions 10.13 Answers.
You have learnt that among seed bearing plants, herbaceous annuals attain a limited height and do not need to increase in girth. Primary growth is often sufficient to meet their structure needs.' However, in woody perennials that reach enormous height and produce large canopies, increase in girth is necessary to support the weight of the shoot. Secondary growth, derived from secondary or lateral meristems results in increase in diameter of stems and roots.
After reading this unit you will be able to: explain the phenomenon of secondary growth in woody plants, describe the structure and give function of each of the following * vascular cambia, * cork cambium, * lenticels, identify the secondary growth of monocotyledonous and dicotyledonous stems distinguish between types of wood, its annual rings, sapwoo$, heartwood and bark, explain the commercial uses of cork and different wood, list various types of unusual secondary growth in stems and roots.
10.2 SECONDARY GROWTH IN A TYPICAL DICOTYLEDONOUS STEM
You have already read in unit 8 that the primary plant body is in itself structurally and functionally complete; for example the majority of monocotyledons and pteridophytes. In gymnosperms and most dicotyledons primary growth is followed by secondary growth. In stem, the secondary growth in thickness in diameter is confined both intrastelar, i.e, within the.stele and extrastelar regions. The cells that form secondary tissues are produced by lateral meristems. The lateral meristemsgrow and join to make a circular ring known as the vascular cambium which lays down cells that become the secondary vascular tissues. In the stem, cells which are situated between the primary xylem and primary phloem in the, vascular bundles become meristematic and form part of the vascular cambium. Additional cells between the vascular bundles also become meristmatic. Hence the vascular cambium can be seen in a cross section of the stem as a continuous ring of tissue, wih h e xylem and pith on h e inside and phloem, cortex, and epidermis on h e outside
Fig. 10.1: Diagrammatic representation of secondary growth in a dicot stem upto two yean (stages in T.S.)
The vascular cambium usually, if not always, has a dual origin within the primary tissues; from provascular strands, and from the "ground" meristem tissues between those strands. These two modes of origin are termed intrafascicular (within fascicles) and interfascicular (between fascicles). The term "provascular tissue" will be used in the text. We should know what this means. Provascular tissue is the precursor of all vascular tissues and "Procambium" is that part of the provascular tissue that is the precwsor of the vascular cambium (which may also produce some metaxylem). The transitional stages between procambium and cambium are denoted as metacambium. Both procambium and then metacambium differentiate acropetally within the provascular bundles. If we see the transections of Populus, metacambium can be first detected by a series of radial, anticlinal divisions in laterally extended sets of tangentially aligned cells. Most of the divisions in this layer are periclinal, producing metaxylem and metaphloem. The cells between the metaxylem and metaphloem eventually begin to function as cambial initials. Cambial initials consists of two m'orphological types of cells-axially short, blocky, ray cells. Procambium at first consists of short cells from which longer cells may arise in two ways (Soh 1990): (1) Different cell lengths result from locally different rates of transverse andlor pseudotransverse cell divisions during growth. Thus the shorter cells become ray initials and the longer become fusifonn initials (2)All procambial cells first become quite elongated. Then some of them.by nonrandom transverse and/or pseudotransverse divisions are secondarily transformed into sets of axially short ray initials. As soon as a circle of vascular cambium is completed, its cells divide ta produce
Residual procambium a1
Fasc~cularcarnb~um Interfaslcular camblum Prlmary xylem
Primary phloem Primary phloem
Fig. 10.2: Formation of complete vascular cambium. A. After completion of primary growth some meristematic cells remain between primary xylem and primary phloem. B The . residual procambium become reactivated to form fascicular cambium and some parenchyma cells of pith become merktematic to form the interfascicular cambium. C. Formation of complete cylinder of vascular cambium. D. Cylinders of secondary phloem and secondary xylem have been formed by vascular cambium.
is differentiated. In the mean time cells produced just outside the vascular cambium becomes differentiated into secondary phloem. Thc or~gin the of . the main axis of the cambium is continuous with that of the branches. It divides and produces new cell towards the outside.The tasks of absorbing water and minerals are performed by younger roots at the far ends of the root system. In almost all the dicotyledons and gymnosperms. including monocotyledons. which originates from interfascicular parenchyma. the occurrence of apical growth. the first formed secondary phloem is destroyed and replaced by the newly formed secondary phloem. As the cambium produces new wood. The procambium and cambium may be looked upon as two developmental stages of the same meristem. 10. Define secondaky growth.3 VASCULAR CAMBIUM-GENERAL DEVELOPMENT AND STRUCTURE ' i I 1 t ! In certain plants. These cells become suberised i. Simultaneously another lateral meristem called phellogen (formerly termed cork cambium). Commonly the bands of fascicular cambium become interconnected by additional bands of meristem. because it originates within the bundles or larger segments of the primary vascular system. all the cells of the procambium undergo differentiation into primary vascular tissues. Name the region in a dicot stem where secondary growth occurs. The epidernlis is replaced by a new protective layer of secondary tissue.new cells. the stem grows thicker with secondary xylem and phloem. A detailed account of the structure and description of secondary tissues and secondary growth will be given later in the unit. extending through the nodes and internodes.lO. the peripheral primary tissues. a portion of the procambium remains meristematic even after the completion of primary growth and develops into the cambium of the secondary body. The main roots of a tree are large and woody and provide anchorage to the plant. b. (Fig. Those formed inside the ring of cambium differentiate into secondary xylem or wood. impregnated with a waterproof waxy material and die. the phloem peripheral to the vascular camhium becomes stretched.. giving rise to a protective layer of cork. the interfascicular cambium. A completely formed cambium of the stem has the shape of hollow cylinder. and the precise method of division in a tangential plane during the formation of xylem and phloem. they intergrade with regard to their morphological and physiological characteristics.2) With the addition of secondary tissues. Most of the cells of secondary xylem have very thick walls.As more secondary xylem is formed. and participate in the transport of organic substances. Secondary Growth SAQ 1: a. and it may extend some distance into the leaves. The cambium that arises within the bundles of primary vascular tissue of the stem is called fascicular cambium.e. the stem increases in diameter. cortex and epidermis become compressed and destroyed. about which you would study in the later part of this unit. The typical features of cambium of a woody dicotyledons and gymnosperms is segregation of its initials into fusiform and ray initials. Secondary growth in roots is similar to that in the stem. If the axis is branched.
. (Bailey.. It is from this point that the cambium produces the secondary xylem centripetally and secondary phloem centrifugally. cell phloem differen tdates from this cell Seconu xylem layer differentiates surface of stem or root .3. . no cytological changes are noticeable in connection with the return to meristematic activity. . they have 14 contact faces with other such cells. cambium is divided into: i) Storied or Stratified Cambium: The groups of ray initials may become taller either by the loss of fusifonn initials located between two groups of ray initials. Fusiform initials sometimes become very long in old trunks of Sequoia sempewirens e.3. 10. . . 10.. vascular cambium cell xylem differentiates from this . 10.lO. convert itself into a row of ray initials..Plant Development-11 interfascicular cambium in the more or less vacuolated interfascicular parenchyma results from a resumption of meristematic activity by a potentially meristematic tissue. Fig.. In most of the dicotyledons and gymnosperms the cambial cylinder develops between the priillary xylem and phloem..3: Production of new xylem and new phloem each year. Usually. All the structural elements which extend radially are produced by the ray initials... allowing them to fuse..1 Structure of the Vascular Cambium: In gymnosperm initials 1000-8700pm long were reported. These cells are shaped somewhat like flat shoe laces. . (Fig.g. they can often have up to 32 faces with an average of1 8. This meristem. a position that is retained throughout the life of the plant.... - . they reach a maximum length of 8700p m.. The mean length of fusifom cells varies. ii) Ray Initials: They are smaller than fusifom initials and are nearly isodiameteric (equal dimensions...2 Types of Cambium: On the basis of the arrangement of the fusiform cells as seen in tangential section.4 A).. It would be easy to assume that they have about 8 faces but a study of Pinus sylveslris has shown that 8 sides are minimum. . . 1923).. among taxa and within an individual plant.3). Vascular cambium is shifted away from the centre of the plant....(Fig.. ... :: :.. . small slides) or only about two or three times as tall as wide. ii) Ray Initials i) Fusiform Initials: These are typically axially elongated cells with tapered ends. or a fusifom~ initials can by transverse division. It tends to increase withthe age of the plant.. 10. in the anatomical sense of the tern1 usually includes two histologically distinct kinds of cells: i) Fusiform Initials. Now we will study about the strucuture of vascular cambium..
Storied cambium B. In a cambial fusiform cell. They are obviously. and the cambial cylinder also enlarges in circumference. Conversely. They also differ cytologically from the other meristematic cells.3. (Fig. 10.4B) Secondary Growth Stoned cambia occur in about 50 families of dicots. i' I I The thickness of xylem cylinder increases by secondary growth. views of fusiform initials and ray initials A.In this type the fusiform cells are arranged in tiers.4: L.3 Structure of Protoplast and Cell Growth: Active cambial cells are richly cytoplasmic and are not wholly undifferentiated. different in shape from embryonic ground meristem and apical meristem cells. although differentiation of camhid derivatives ! . the xylem cells towards the interior of axis and the phloem cells towards its periphery. 10. Fusiform lnitials -Ray Initials Fig. or stories. That is. ii). Thus nonstoried initials are longer. Storied cambia do not occur in gymnosperms. but commonly not in all genera of a family. Such radial seriation may persist in the developing xylem and phloem or it may be disturbed through various kinds of growth readjustments during the differentiation of these tissues. In nonstoried cambia there is no lateral alignment. Although active cambial cells undergo repeated periclinal divisions and radial growth. Active fusiform cells commonly have one or hvo large vacuoles transversed by many slender cytoplamic strands. Non-storied or Nonstratified Cambium : In this type of cambium the ends of cambial fusifoim cells typically overlap much more extensively and in a seemingly random manner. The consistent tangential orientation of the planes of division during the formation of vascular tissues determines the arrangement of cambial derivatives in radial rows. have large mitochondria and often have more highly differentiated plastids (Catesson 1990). They are more highly vacuolated. If you view them tangentially the ends of cells in axially adjacent stories generally overlap only slightly making a zigzag pattern. Non-storied cambium. the ends of large tangential groups of cells are aligned at the same levels of axis.S. Whereas in a ray cell it is usually more nearly spherically. the width of cambial zone does not increase indefinitely. and small vacuoles in the peripheral cytoplasm. the nucleus is quite elongated. The cambium initials form pholem and xylem by tangential division. In vesselless dicotledons the fusiform initials may reach a maximum length of 6200 pn . They are also of more common occurance. These vascular tissues are laid down in hvo opposite directions.lO.
Towards (i)Whole sell Thngcntisd Divisnan (iii) Ncrt (E) Tangential Division ss in TS (i) Pmnehypra cell af msdullw ray ' (ii) First tanpntlsl wall . However. The rate of production of cambial derivatives depends on the number of cells in the cambial zone and on the duration of the cell cycle. Addition of new fusiform initials is brought about by longitudinal anticlinal divisions of the existing initials in the storied cambium while in nonstoried cambium. there are togelher referred to as c ~ m b ~ rone al Fig. Let us now study how the ray initials are formed in the cambium. followed by intrusive growth. the balance is often imprecise.3. Ray initial Because of the excessive length of the fusiform initials. 10. The plate takes a long time to reach the end walls. Fusifonn cambial initial .5). the zone itself does not disappear. and each of the new cells becomes as long as or even longer than the cell from which it was derived.6). the d o l e v einto xylem & D V ~ S S ~ rzone Remaintmg thln ~ l l c d ss Xylem Cell (a) rernmoa msnrrtemat8q new cells a' aod a' Imk rimtlar to (a). 10. the fusiform initials undergo oblique. iv) In the last whole or part of a fusiform initial may be segmented by transverse divisions to form a tier of ray initials. ii) A single cell may be cut off the end of a fusiform initials. iii) A declining fusiform initial may be reduced to a single ray initial.Plant Development-11 into xylem and phloem continually removes cells from the cambial zone. The cell plate begins to form between the two nuclei and it spreads slowly. 10. A.4 Ray Initiation: With the enlargement of the cmbial cylinder new r& intials develop and single fusiform initials are continuously lost from the cambium and are replaced by new ones (Fig 10. B. Second tan~cntialwall . i) A single cell may be cut off along the side of a fusiform initial (lateral division). and cambial zone thickness tends to vary during the active season. the formation of the cell plate during the process of longitudinal division is peculiar to these cells. (Fig.5: Cambial development. If the rates of radial growth and periclinal division are just balanced by the rate of cell loss through differentiation. But various degrees of transverse divisions may occur between these types. . pseudotransverse anticlinal divisions. the cambial zone thickness remains constant.
........ A-Division of fusiform cambial initial to uniseriate....... B-Multkriate ray. b.... ~ifferentiak between Storied or stratified cambium and Non storied or nonstratified cambium. c.............Secondary Growth Ray.. many Cells in TS Fig........... . Describe the types of cellsfound in the vascular cambium..........6: Origin of a secondary ray... 10.. SAQ2: a. Name the two cambia which join and form a cylinder of of cambium.... ...
1924 the tree is growing straight again. In geographical regions where there are clear cut seasons. However. 1930 A fire sweeps through the forest. making it lean. the rings are now wider on the lower side. But it neighbours are growing too. an extensive tubular structure having the general form of the axis of the plant. 10. This results in a wide cambial zone. There are some plants in which cambium is active during the entire life. and the~r crowns and root systems take much of the water and sunshine the tree needs. These plants commonly occur in the tropical regions where seasons are mot markedly different. and year by year more and more of the scare is covered over by newly Formed wood. . therefore.X T S ofdicotyledonous stem showing annual rings.Plant Development-I1 - 10. Thcse narrow rings may have been caused by a prlonged dry spell.7). By analysing the tree rings.4 CAMBIAL ACTIVITY - The radial growth is directly correlated with the rate of cambial activity. 10. 1927 ' The surrounding tree an harvested. 1994 When the tree was 6 ycars old. each one of which represents a seasonal growth. 1942. these layers appear as rings. This study is known as Dendroclimatology (See box 10. and the terms annual ring and growth ring or growth layer are applied to each layer. The tree can now grow rapidly again. . An annual ring or growth ring of xylem is a layer of secondary xylem fornled in one growing season over the entire plant and is.1 Formation of Annual Rings: The secondary xylem in perennial axis commonly consists of concentric layers. cambium shows decreased activity with the onset of autumn. something pushed against it. 1957 -nother series of narrow rings may have been caused by an insect like the larva of the sawfly. It is open at ends where meristems occur. The wood thus formed in spring is called springsummer wood or earlywood and that formed in autumn is called autumn wood or latewood.. at the time of cambial activity cell divisions are faster than cell differentiation. 10. This mriy last till the beginning of the following spring. If you see a cross section of the axis.1). as the tree builds "reaction wood" to help support it. and it enters a dormant state during winter. In warm temperates climates. a great deal can be learned about past climatic conditions. One or two dry summers would not dried the ground enough to slow the Iree's growth this mueh. These variations result from internal and external factors. the tree is only scarred. But as soon as differentiation begins. The larger trees are removed and there is onee again ample nourisment and sunlight. Fig. The vascular cambium shows variation in the period and intensity of activity. The variation in the number of cell across the cambial zone seems to express the balance between the rate of cell division and the rate of differentiation of the derivatives. It eats the leaces and leafbuds of many kinds of coniferous trees. the percentage of ringless trees is still lower. However. a balance is established and the width of the zone remains more or less constant. When the rate of division becomes less and the rate of differentiation faster the cambial zone becomes narrower. Thus periodical activity of cambium results in the formation of ring wood. In spring the cambial activity is resumed and becomes maximum during summer.4. not all tropical trees exhibit a continuous cambial activity. It is noted that each annual ring corresponds to one year's growth hence the age of plant can be determined roughly by counting the total number of-annual rings in a log (trunk) as seen in a transverse section (Fig. Tree ring analysis is also known as Dendrochronology. Fortunately.
if a certain year is drought year then in that particular year much smaller wood layer's will be produced. the size of each ring varies depending on environmental conditions. 10.BOX 10. This will result in a marked decrease in photosynthesis. Then by matching the rings one can know the exact age of the living tree or trees (Fig. including precipitation and temperature. Then similar patterns appear in the rings of many tree species in a large geographical area. For example. In spring growth is fast and wood is light in colour. Some times the variation in tree rings can be due to a single environmental factor.8). causing low wood production. For example. In addition other useful information can be obtained by analysing tree rings as weII. in trees that have lived for several thousand years first a master chronology of complete records of sample of rings dating back as far as possible is developed.8: Tree ring dating: A master chronology is developed using progressively older pieces of wood from the same geographical area The age of the sample can be accurately de termined by matching the rings of a wood sample of unknown age to the master chronolop . This will also result in two annual rings being very close each other because very little gqowth wiIl take place. Some times locusts may have eaten the leaves just after they have appeared. To study the sequence of rings. 10. In summer the wood is darker.1: TREE RING ANALYSIS -A WAY TO TELL THE LIFE HISTORY OF A TREE Secondary Growth Every year a new growth ring is added to trunk of the tree. Tree ring dating Fig. By counting the dark rings you can tell the age of tree.
The axial . phloem and cortex. The horizontal system is made up of xylem rays (Fig.5 SECONDARY XYLEM -- The products of the camb~um formed towards the centre of the stem and root constitute secondary xylem. xylem ray cclls and sometimes secretory cells.9) and the vertical or axial system consists tracheary elements.1 Basic Structure of Secondary Xylem: Secondary xylem is characterised by the existence of two systems of elements which differ in the orientation of their longitudinal axis. parenchyma cells. 10. 10. The rings are examined counted from the core and then analysed. To determine the age of living old tree you do not have to cut it down. to examine the rings.and the arrangement of these elements vary in different group of plants. The hole is then treated with a disinfectant and covered up without harm to the tree. different types of fibres. It is driven in the stem of a tree and a core of wood is removed. make it possible to identify individual specles on the basis of secondary xylem alone. Secondary xylem is composed of tracheids. USA studied ring widths in trees from various sites. The quantitative differences in the number of cells and the size of the elemcnts that exist between the species of a single genus. a) b) b) All. whereas fusiform initials from axial parenchyma. Douglass.5. Regions with warm climate have a low per cent of ringless trees. They found that a very significant statistical relationship exists between the growth of trees and climatic condition.2 Wood Parenchyma: Two types of parepchyn~a found in secondary xylem : The axial parenchyma are and the ray parenchyma. special cambial initials give rise to ray parenchyma.E. Write T for true and F for false in the given boxes.5. which in turn is connected with the living cells of the pith. The relatively short. Prcsently by ring analysis data are gathered to determine climates of prehistoric times. • • It is possible to calculate the approximate age of a tree by counting the total number of rings in one log of wood. With the help of computers and statistical analysis they developed techniques that takes into account of climate and other environmental variables. The increment borer is primarily made up of a rigid metal cylinder. The occurrence. The living cells of the rays and of the vertical system are usually interconnected and a continuous system of living cells is formed. Fritts and others associated with the Laboratory of Tree Ring Research of the University of Arizona Phoenix. For several years A. A simple instrument known as increment borer is used. SAQ 3: Which of the following statemms are true. One system is horizontal and other vertical. fibres and wood parenchyma. They were able to reconstruct relatively precise histories'of climatic changes and fluctuations dating back thousand of years. 10. Harlod C. vessel members.Plant Development-11 A great deal can also be studied from tree rings about the past climatic conditions. 10.tropical trees cxhibit a continuous cambial activity.
. ii) in which it is vertical. of The ray parenchyma may be of different kinds. 10.CROSS SECTION Xylem ray Secondary Growth tracheid lyma cell 1 Resin duct TANGENTIAL SECTION Fig. the wr.10). 10.9: Structure of secondary xylem in three dimenshnal diagram of a cube ofPinus. I i I parenchyma cells may be as long as the fusiform initials or much shorter. 10. End wall Pits -Pits Parenchyma -LTracheids Tracheids Fig.10: Basic structu~e xylem. It is more common to find shorter parenchyma cells (Fig. End wall i . most common forms are : i) in those which longest axis of the cell is radial.
Boarded pit. The oarenchvma. The events that occur in the formation of heart wood include disintegration of protoplast. it is multiseriate. . 10. -----.The nucleus and " .. The outer part of secondary xylem with living parenchyma is named as Sapwood or alburnum (Fig. In nearly all the trees the central portion of the xylem consists of dead parenchyma which ceases to conduct water. r i 1 In several plants wood parenchyma cells form protuberances which penetrate into the vessels through the pits after they become inactive. silica bodies and other substances are also deposited in these cells. tanins. resins. 10. width and height of each ray can be measured by cross sections and tangential sections respectively. gums. and . part of cytoplasm of the parenchyma cells may enter the tyloses. --cells of the xylem serve to store reserve food materials such as starch and fats. B.its (Fie.. when two cell wide it is biseriate and when more than two cells wide. Tyfoses may also divide. These secondary cell walls are commonly characterized by the presence of depressions or cavities varying in size.10. The ray parenchyma is the main route of radial symplasmic transport between xylem and phloem. Oil. aromatic compounds and coloured substances which develop in the cells are accumulated in the heartwood. The amount of heartwood and sapwood varies in different speices. 10. Resin duct in Pinus wood. halfbordered. The outer part of the secondmy xylem contains living cells and at least one or two 6utermost rings participate in the conduction of water. This is called heartwood or duramen. These differences are influenced by genetic and environmental conditions. When the ray is one cell wide it is called uniseriate ray.11: A. These outgrowths are termed tvloses (singular : &lose) .11 B). All the cells of'ray parenchyma may have primary wall or only secondary walls are found. 10. d e ~ t h structure. Although the formation of tyloses is considered a natural phenomenon. in many species it has been reported to result from mechanical injury or diseases.5. .The length. or later injured. .3. Heartwood and Sapwood: Heartwood may sometimes develop as the result of pathological conditions. In such species in which the tyloses are formed the inner portion of the cell is totally blocked by the tyloses.12). Tannin. and sometimes even bordered. crystals. Such cavities are termed . loss of cell sap and hydrolysis of reserve material stored and formation of tyloses. The secondary walls may develop pitpairs that are to be simple.Plant Development-11 The number of xylem rays increases with the expansion of stem girth.11 A). - Secondary wall - Border Torus ---- Primary wall 7 Globule of resin Secretory cells \ Fig. (Fin.
when the lumen is small the wood is dense and heavy. resin and oils largely determine the durability of wood. Some other important uses of Wood: Teak. We will discuss some of the properties of wood by which wood quality is judged. i) Weight: The wood may be either light or heavy. bridges. structural works. The majority of well known commercially important wood range from 0. Strong woods are used for building. ships. carts. Thus the dense and heavy woods are of greater strength. The durable woods are used for ship building. 10. railway wagons. wagons. Wood has much importance in day to day life in form of furniture.65 in specific gravity.35 to 0. railway sleepers. In the timber trade wood of dicotyledons is known as hardwood and that of gymnosperm as softwood. paper gum. resin and several other industrial purposes. boats. is used extensively in insulation. railway sleepers and furniture etc. thick walled fibres make wood heavy. bodies of campartments.12 to 0. furniture etc. Extremely light woods such as Ochroma is also found.35.12: Heartwood and sapwood in C. ii) If a large proportion of the wood is made up of fibers or fiber tracheids it tends to be strong. Balsa wood (Ochroma lagopus) and sola (Acshynomene) have abundant of parenchyma and very few fibres. However. If we see the histological structure of the wood of dicotyledon and that of gymnosperm there are . Balsa (Ochroma) with a specific gravity of 0. there is an important difference in the wood of dictoyledon and gymnosperm.5.S. Light woods have limited economic importance. The abundance of slender. Both light and heavywoods can be durable. The former have vessels and the later lack them. sisham and rose woodware decorative timbers for panelling furniture. railway compartments. masts.4 Economic Importance of Woods and its Characterjstics. The presence of tyloses and other natural constituents of wood such as tannin. The heavy woods are used in construction.periderm I"cnrk"l Phloem I I Heartwood A Sapwood Fig. These terms do not accurately express the degree of hardness.fundamental differences. Differences in weight are due to variations in the proportions of wall substance and of lumen space . as in both groups wood with both hard and soft structure can be noted. of mulberry trees. . 10. as material for modelling (by hitects) and for life rafts. iii) Durability: The ability of wood to withstand decay by the action of fungi and bacteria is largely dependent upon the chemical nature of the wood and is referred as its durability. construction. carts.
.. What structural features gives strength to wood? ... . c........ matches........ music instruments...Plant Devela cabinet... List as many independent uses of wood as you can think............. pattern making in cabinet........ Distinguish between hardwood and softwood......... cosmetics....6 SECONDARY PHLOEM In the secondary phloem also there are two systems the vertical and horizontalproducts from those cambial intials as in the case of *!em.... How do uniseriate... ..... 10..... roxburghii is used for house building.. biseriate and multiseriate rays differ from one another.... soaps.. e.... for carving idols.... boxes and matches. boxes.... b... d.. aspects of art etc... SAQ 4: a.......... In India P.......... v) Sandal Wood: Sandal wood is smooth and tends itself to exquisite used for carving as well as for extraction of the oil.. Though these two tissues secondary xylem and secondary phloem differ in olttageny and structure at maturity....... windows... railway sleepers etc.. packing cases..... The sandalwood oil is fragrant and finds use in perfumes... incense sticks etc. List the two types of systems @d various cells found in secondary xylem.... inlay works. iv) Pines: They are chiefly used for doors.
in Tilia the sieve tubes are active for several years including winter.6. The phloem may contain well developed . whether it is storied or not. As you have already studied.lO. 10. Thus xylem and phloem rays are continuous. This is possibly related to its role in protecting the plant. Plasmodesma Phloem Sie\e tube cell Phloen~ cells Fig 10.1 Economic Value of Secondary Phloem: The secondary phloem may be quite rich in secretory tissues. As in the xylem. Secondly it depends upon the extent of elongation of various elements of vertical system during the differentiation of the cells. that the ray initials in the cambium produce cells towards both the xylem and phloem. This is because cells which are produced at the beginning of the season are extended radially while in the end of the season they are flattened. In the dicotyledons the functional secondary phloem is restricted generally to that produced in the last growing season. growth rings can be observed in the secondary phloem but they are less distinct than in secondary xylem. The horizontal system consists axial and ray phloem which is made up of parenchymal cells.e. The widening of the phloem ray may be accomplished solely by the lateral expansion of the existing cells or as is more common by an increase in the number of cells on the periphery by radial divisions. The primary reason is also because of absence of lignin that is why they can not be used as an indication of the age of secondary phloem.The important components of the vertical system are sieve elements. the arrangement of the tissue in phloem is primarily determined by the nature of cambium i.13). However. In some cases when the cambium starts producing new phloem. phloem parenchyma and phloem fibers (Fig.13: Basic structure of phloem I L I r In several species of dicotyledonous trees. After some growth the arrangement of growth ring becomes obsecured due to the obliteration of the sieve elements. In the vicinity of the cambium the xylem and phloem rays are equal in size but in many plants the mature outer portions of phloem rays are wider. almost all the previously produced sieve tubes cease to function.
the increase in thickness is through secondary cambium (which is entirely a secondary menstem). A phellogen produces phelloderm adaxially and phellem abaxially (Fig.duct systems in some species. forming a protective tissue with suberisation (Dracaena) are present. c..14). sunnhemp etc. cambium is absent. the cambium. They do not produce secondary phloem or xylem outside and inside respectively as in dicotyledonous stems and roots. a. Thus the vascular bundles remains embedded in the conjunctive tissue. SAQ 5: Write T i'n front of true statement and F for false statement in the bracket provided. These newly formed inner secondary tissues directly differentiate into oval shaped collateral vascular bundles and radially arranged parenchyma cells called conjunctive tissue. . They are rather easy to separate from underlying woody tissues. similar to vascular cambium. But in some herbaceous and treelike woody monocotyledons plants belonging to families Liliaceae. Important sources of bast fibres are flax. The cork tissue forms a protective layer of the tree after the epidermis dies and is shed. 10. In a monocot stem a periderm is absent but some storied cork cells. ( In several dicotyledonous species we can see more distinct ring in secondary ) phloem than in secondary xylem. ( 10. It is the system of lactifers in the bark of Hevea that is tapped to obtain rubber and the resin canals of the bark of conifers that is are harvested for pine resin which is further distilled to make turpentine and resins. a few layers in thickness. ( ) We can judge the precise age of tree by counting the number of rings of ) phloem. The secondary cambium is made up of rectangular fusiform cells. these tissues senescence and die. the cells divide tangentially forming a band of secondary . Bast Fibers: These are selerenchyma fibers associated with the phloem of certain stems of plants.7 SECONDARY GROWTH IN MONOCOT STEM Normally in a monocot stem. Cork is generally formed in the stem and root of dicotyledons which have a continuous and pronounced secondary thickening cork is not formed in leaves with the exception scales of winter buds of certain plants.e. This meristem. jute. is generally suberized and inhibits translocation of water and solutes to tissue abaxial to it. no secondary growth takes place. As a result. as the vascular bundles are closed i. the phellogen. In phloem there are two systems the vertical and horizontal. Agavaceae etc. Instead the secondary cambium cuts of secondary tissues on the inner side first and then a small amount of new tissues on the outerside. cambium. is a uniseriate layer of initial cells that by periclinal divisions gives off derivatives from their adaxial and abaxial faces. b. Before dealing with periderm formation we must know what periderm is? The tenn periderm is collectively given to the protective tissues phellem and phelloderm and the meristem that lies between them and gives rise to them. At the time of secondary growth some of the innermost parenchymatous cell become meristematic. Phellen. They arise with primary tissues from the apical meristem or with secondary tissues produced by the lateral meristem. a major constituent of periderm.
3 10. They are similar to the parenchyma cells of the cortex but. Secondary Growth Epidermis Phellem Phellogen Phelloderm Cortex 10. Fig.14). in some plants two periods of phellogen activity have been noticed in a single annual period of cambial activity. They may contain crystal containing cell.8. and produced centripetally by the phellogen. In some plants cdls of the phelloderm have chloroplasts and are photosynthetic. The cells of phellem divide tangentially. The cells are uniform.8.2 Phellem Phellem or cork arises from the abaxial derivatives of phellogen.Jn a cross section cells are devoid of intercellular spaces except in lenticular region. 10. The phelloderm-a parenchymatous tissue in some species. In certain species the cork cells's primary walls are suberized and contain a thick suberin layer interior to the primary wall called the suberin lamella. rectangular in cross section with their shorter axis in the radial direction. if the phelloderm is multiseriate. The activity period may or may not concide with that of the cambium. Peridem is usually divided into three parts: i) ii). This phenomenon of impregnation of walls with suberin is referred as suberization.14: Periderm formation. they are usually arranged in radial rows. Cork cells are dead. and radially flattened and divide tangentially. The phellogen has distinct periods of activity and nonactivity. where they occur in alternating layers in Betula this feature causes an interesting feature and cork is peeled off like sheets of papers. 10. However. iii) The phellogen-cork cambium The phellem-cork which is produced centrifugally by the phellogen.S they appear as regular polygons. Of Of cork The may found in one species e. Arbutus and Betula. This substance suberin is highly impervious to gases.1 Structure Phellogen is a lateral meristem consisting of a single layer of initial cells.g. .8.3 Phelloderm: The phelloderm cells are Kving cells with nonsuberised walls.Cork is an important part of secondary tissue which is termed periderm (Fig. In a T L. The cells of phellem or cork are usually polygonal. water and resist the action of acid. or even nonsuberised. The protoplasts of phellogen cells contain vacoules of various sizes and may contain chloroplasts and tannins. they may be sclerieds. Periderm 10. Phellogen has no intercellular spaces except in the regions of lenticels.
collenchyma or a parenchyma cell. 10. Phellogen Phelloderm A Phellem Phellogen Phelloderm " B Fig. of which the inner ceases to divide further. Quercus suber which occur naturally in the . The stripped cork shows a rough outer surface and a smooth inner surface. If the firstformed periderm remains on the axial organ for many years. the other layers of cork show cracks and are shed off. It may remain on the plant for an indefinite period of time.Plant Development-I1 10. The outer divides periclinally.15: Formation of periderm stages. The outer cells differentiates into the cork cell and the inner constitutes the phellogen initial and continues to divide. 10.5 Commercial Cork The source of commercial cork is. In this species phellogen arises in the epidermis. The number of phellem layers is usually greater than the number of phelloderm layers.Epidermis . The subsequent phellogen produces cork more rapidly and in about 10 years a sufficient thickness to be of commercial value.Cuticle .4 Origin and Development of Periderm The phellogen may differentiate in a living epidermal (Fig. to keep pace with the increase in the circumference of the cork cylinder. . Thus the thickness of cork on a plant remains constant.8. hvo similar cells are formed. lS). . Just before the onset of meristmatic activity the cell loses the central vacuoles. Following the first periclinal division. For commercial purpose the first formed periderm is removed when the tree is about 20 years old and about 40 cm in diameter. The expased cells of the phelloderm and cortex dry out and die. The initials of phellogen cells occasionally undergo anticlinal divisions.10. the volume of cytoplasm increases and it undergoes a periclinal division.. and a new phellogen is formed a few millimeters within the cortex. countries boarding the medeterranean sea.8.
Therefore. Here the lenticels form collar like structure around the thinner roots. Histologically phellogen iii) i s impermeable to water gases and can withstand the action of acid.16). Large corks are usually cut from sheets of ground and compressed cork or from "multiple sheets" composed of layers cemented together. iv) alternating periods of activity and inactivity.There is continuity of intercellular spaces of lenticels with tissue in the axial organs. Due to many intercellular spaces. Stoma Complementary cells Phellogen Fig. Cork is several centimeters thick and the lenticels remain active for a long time and result in the formation of cylinders of complementary tissue which extend from the phellogen to the surface of the phellem. b. lenticels have a loose structure.The cork is valuable because it is impervious to gases and liquid. lenticel protrude above the periderm because of their larger size and loose arrangement of numerous cells. Cork is unparallel as a material for making stoppers for wine and champange bottles. Periderm i s usually divided into I n some plants phellogen has Section B i) consists only one type of initial cells.16: Lenticel in L. They are usually found on stem and roots and appear on young branches or other organs as rough dark patches. Sheets of cork from the tree are rarely more than 3 cm. It is used for insulation. You may like to know why commercial cork is to be cut in a particular plane. 10. d. These kinds of cork are of low quality.9 DISTRIBUTION OF LENTICELS Lenticels are highly differentiated lens shaped areas of periderm. ii) three parts a) Phellogen b) Phellem c) Phelloderm c. Cork with a diameter greater than that can not be obtained by cutting in the usual manner. This complementary tissue forms the patches of dark brown crumbling tissue found in commercial cork. I n the roots ofPhoenix dactylyeera lenticel like structure occur which take part in aeration of the root but which differ from the described ordinary lenticels. bottle corks to be in a direction parellel to the surface of the trunk. . early stage in fo~matian. elasticity and lightness. Mostly scattered over the entire surface of stem (Fig. Cork 10. Secondary Growth SAQ 6: Complete the sentences from section A with those given in section B: Section A a. Because of the radial orientation of the tissues. 10. nonreactive and has strength.S. lenticels are believed to participate in gas exchange. in the periderm. You must also know about lenticels: These are restricted area of relatively loosely arranged cells. This way the cylinderical lenticels extend transversely through them. sound proofing and in the manufacture of sportsgoods. thick.
10. Campsis radicans Vitis and some other swcies. The first formed lenticels generally appear below a stoma or group of stomata. 10. Lenticular phellogen has more. .S) -well formed In the temperate regions lenticels become closed by the end of autumn season by a closing layer. Whereas in some plants lenticels are formed relatively early in the life of the plant and are shed together with the bark. Various terms such as cambial variants. many ofwhich areclimbers do not possess IenticeIs. 10. Fig. as well as those produced towards the exterior by the phellogen of the lenticels are termed complementary cells. The cells which are derived from the divisions of the substomatal cells.Plant Development-I1 If we examine lenticel it usualfy looks like a convex lens both internally and externally. Cambium shows variation in its activity giving rise to conditions which are rather typical. in others they may remain active for several years. Cells below the stomata begin to divide in different directions and the chlorophyll in them 1 disappear so that a loose colourless tissue is formed.17: Lenticel (in L.g. SAQ 7: Define the following in two or three sentences: Lenticels and Complementary cells.1 Development and Structure of Lenticels Lenticels originate form localized regions in the phellogen that become continuous with the nonlenticular phellogen. Philadephus Anabasis. Haloxylon.9.10 CAMBIAL VARIANTS In this unit you have studied the function of vascular cambium and cork cambium in dicotyledonous and monocotyledonous stems accounting for development of secondary tissues.intercellular spaces and produces derivatives at a higher state than do nonlenticular phellogen. 10.17). The cells that are derived from the divisions become more and more periclinal until the phellogen of the lenticels is formed (Fig. After division the number of cells increases and masses of complementary cells are pushed out and rise above the surface of the organ. anomalous or . Complementary cells Only a few plants e.
10.18). n C. .) variant has been recommended for usage. Instead cambial Secondary Growth Fig. ' l Serjania.aberrant secondary growth have been used to describe these instances (Fig.19: T S stem Bignonia i. Epidermis Parenchyma Sclerenchyma Cambium Strips Starch Sheath Pith Phloem Xylem Fg 10. anatomists have discouraged the use of the term anomalous secondary growth.1 I Stems n I. 73 . D. In Bignonia (Fig. I n Bauhinia. showing cambial variant. 10.18: Cambial Variants (Anomalous stem structures. phloem is much more abundant than xylem.10. 10. sp.. As the variants are of quite regular occurence in certain plants. Thus in some part of the plant the amount of xylem is much greater than phloem while in the other. Its products show unusual arrangement and proportion. A. The cambium produces secondary xylem and secondary phloem in dififerent amounts. 10. This feature results a Phloem can be identified by characteristically ridged and furrowed xylem cyl~nder. I n Boerhaatba. The cambium i s persistent and normal in position. In Bignonia ~ .19) and some other members of the Bignoniaceae family.
There. These groups of bundles function as independent cylinders and give rise to separate cambia. Tinosporaetc. 10. The Cambium is abnormal in position but normal in activities: I This situation is noted in some climbing species of Ser~ania (Fig. broad and long medullary rays and fluted vascular cylinders are formed. acomplete ring of cambium is formed. four such wedges symmetrically arranged and corresponding in position to the larger primary vascular bundles. so that groups of bundles are formed. The fascicular cambium functions normally but the interfascicular cambium Secondary xylem B Primary Xylem I I FiglO.S stern Serjcrnia 74 I .21).Plant Development-11 the presence of wedges. each of i I I I I I I I I I I I I I Fig. One year old B.Aristolochia (Fig.21: T. As a result.20: Aristolochia in T. Thus bundles become contricted from the cylinder. They may be cut off even at the procambial stage. Two years old I produces only raylike parenchyma cells.S A.20). I I I I. as usually. b) In some climbing species of genus Vitis (graphs) Clematis. 10. During development the cylinder of primary vascular bundles become notched at certain points. 10.
10. This accessory cambium cuts off xylem and phloem in an inverse order i. The bundles are of secondary nature but their cambial activity soon ceases.22: T. each of which has its own periderm. Just outside the bundles a new secondary cambium arises in the pericycle. Secondary Cmwtl c) Epidermis Parenchyma Starch sheath. At later stage an accessory secondary cambium arises in two arcs on the inner side of the normal wood or towards the pith.S. The bundles formed in this way may be arranged irregularly or in defenite concentric rings In Tecoma sp. This newly formed phloem is intraxylary phloem and is secondary in origin. Thus the stem appears as if it is made up of several discrete woody cylinders.which functions normally and independently. 10. In some species the cambium forms tissues centripetally. consisting of bundles embedded in nonvascular tissue. secondary xylem and phloem are produced in the beginning by the activity of a normal cambium ring.22). a) Anomaly d u e to t h e formation of accessory cambium and its activity: In some species of Chenopodium and members of the Amaranthaceae. the anomalous secondary growth results from accessory cambia.e. A hollow cylinder of vascular tissue or a ring of irregularly arranged bundles. 111. Sometimes the woody cylinder is only lobed. xylem towards the periphery and phloem on the inside. And the secondary xylem merges gradually with the previously formed secondary xylem (Fig. of stem Tecoma showing accessory cambia . - Sclerenchyma Phloem Pith Cambium Xy lem Phloem Cambium Xylem Fig.
Salvadora and the families Amaranthaceae and Chenopodiaceae. 10..g. especially those that serve a storage function. Thunbergia. e.2 I Roots n Cambial variants are also found in the roots of some plants.Plant Development-It IV. b) In Chenopodiaceae the successive cambia are seen in the form of long or short arches. Anomaly due to the formation of interxylary (included) phloem as a result of aberrant activity and position of cambium. Frequently the additional cambia in this family forn more or less entire rings. . List various types of cambial variants. 10. In certain plants there are strands of secondary phloem within the second& xylem. each of which cambium produces xylem toward the inside and phloem towards the outside until a new cambium develops from parenchyma cells on the outside of the phloem. Secondary phloem patches are sometimes embedded in secondary xylem in the form of strands. Medullary bundles (Primary) / Fig. They produce irregularly or spirally arranged phloem strands (Fig. Later a series of vascular cambia arise successively outward.23: T. Chenopodium stem. 10. SAQ 8: i). Bougainvillea. in the pericycle or in the inner cortical layers.10. In these plants cambium differentiates outside the primary vascular bundles.S. Those extra patches of secondary phloem present inside the secondary xylem are known as interxylary or included phloem. in Avicennia.23).
The xylem.25). The bundles are themselves largely parenchymatous with a few lignified elements in xylem. Soon its activity ceases. . (Fig. Then from the cells of pericycle and phloem a secondary cambium ring is formed. b). Thus alternate bands of proliferated pericycle and vascular bundles are formed which can be seen as dark coloured and light coloured rings respectively. B.a). Although cambia are in continuous rings they produce separate bundles which are surrounded by conjuctive tissue. In the sweet potato root of Ipomoea batatas (Fig. SAQ 9: Explain how dark cololured and light coloured rings are formed in the beet roots. Sec .Convolvulaceae) :secondary growth is unique.24: A. The bundles are separated from one another by wide radial panels of parenchyma formed due to the activity of newly formed cambium. 10. Thus the phloem appears to be a portion of root that originally differentiated as xylem. T. All the cambial layers continue to function and produce a large amount of storage parenchyma and strands of xylem and phloem. This is followed by the formation of several concentric cambia. Growth layers Storage Parenchyma Secondary Xylem Secondary Phloem cambium Fig10.24). 10. A considerable amount of storage parenchyma is produced in both the directions.S. has an adundant amount of parenchyma Secondary cambia develop in the parenchyma around the individual vessels or vessel groups. . In the beet roots Beta vulgaris first the cambium ring develops near the primary xylem patches which in turn produces secondary xylem towards inside and secondary phloem toward the outside. Magnified part of the root . of root of sugar beet. Periderm of Secondary Xylem . The cambia produce tracheary elements towards the vessels and sieve tubes away from the vessels.
.. The uly wood .. addition of fresh tissues is possible..... active meristematic regions lie between primary xylem and primary phloem from which continued. Cork cambium may originate in successively deeper tissues: epidermis.. The fascicular and interfascicular cambium joins together and form a complete cylinder of vascular cambium. 10....... cortex and phloem.....have most of their secondary tissues arranged in concentric layers.... phellogen differentiates near the surface of the stem.S.-lar cambia........ Woody dicotyledons.. An interfascicular cambium arises from ray parenchyma cells between vast...Lenticels in the bark facilitate gas exchange. so that the stem now increases only in girth.. Fig. Cork cells have suberin in their wall.11 SUMMARY In general the plants having only primary growth are limited in size and longevity.... The most conspicious tissue is wood (secondary xylem)....... After the procambium strand become differentiated into primary vascular tissue.. which makes cell imprevious to gas and 1iquids. This gives rise to phellem and phelloderm..... In many plants... The divisions in cambium are longitudinal........... secondary growth helps the perennial gymnosperms and dicotyledons to increase the diameter and support the height and growth size. These meristematic cells constitute the fascicular cambium...... 10. of root of sweet potato....25: T..Plant Development-I1 ..
Explain why? What is secondary thickening (growth)? Explain the mode of secondary growth in dicotyledons and a monocotyledons. The age of atree and other aspects of its ecological history can be determined by studying histological details of the annual rings. Certain dicotyledonous stem have cambial variants that contribute to unusual secondary growth. Its products showunusual arrangement and proportions. 3. Older wood ceases to function and in gradually accumulated in the centre to form the heartwood or deadcore which becomes plugged with tyloses. 10. The younger. Secondary Growth a Some monocots such as Agave have true secondary growth. Generally an annual ring comprises one year's growth of xylem. iv) The formation of interxylary phloem because of abnormal activity and position of cambium. What are tyloses? How are they formed? 79 . ii) The cambium is abnormal in position but normal in activities. with a type of cambium that produces secondary vascular bundles and parenchyma. will it remain at the same distance from the ground and more up or down in the course of the next 10 years. Cambial variants may arise from the following situations/conditions: i) The cambium is persistent and normal in position. The scientist have shown that only one or two recently formed annual rings or wood are actually involved in the ascent of sap. Beta vulgaris letc. while late wood has smaller vessels and/or a predominance of tracheids. List the similarities and differences in their secondary structures.usually has relatively large vessel elements. If a nail is driven into the trunk of a tfee say at breast height.12 TERMINAL QUESTIONS I. a Cambial variants are also noted in some roots such as Ipomoea batatas. more peripherally located or living wood is called sap wood.. iii) Formation of acessory cambium and its activity.
. 5....... .. ............. How cork is formed? Explain the structure. Why is orikntation of the elements of cork is important in making bottle stoppers? ....?............ properties and uses of commercial cork..... ..Plant Development-I1 4............ 7.. What is periderm? Explain how it divides and the tissue it produces.... Describe some important features of secondary phloem... Suggest some reasons for preferring to heartwood to sapwood as material furniture....
..........................13 ANSWERS Answers SAQ 1: a) b) Secondary growth is growth in diameter due to addition of new tissues due to activities of lateral meristems........ .......... ................................................. ............................................... Secondary growth occurs both in intrastelar amd extrastelar regions............ 10.......................................................................................................................................................... What are lenticels? How they are formed? What are their functions? 9. 10............ .......... ................................................................. Describe the structural anomaly arising as a result of formation of accessory cambium and its activity provide suitable diagram? ...................... Secondary Growth .............. 11.................................................. ............................ Write the names of plants in which interxylary (included) phloem is formed...................... ....................................................... ......... ................................................... .......................................................................................................................... ......................................... ......... Explain briefly the main features of unusual secondary growth in roots.......................................... 8.......... SAQ 2: a) Fascicular cambium and interfascicular cambium..........................
SAQ 4: a) Secondary xylem can be classified in two systems: a) One system is horizontal and b) other is vertical. Biseriate Ray: When two cell wide the ray is called biseriate rays. b) Uniseriate Ray. d) e) i) ii) building boats.3. boxes. vii) packing cases. doors. ships.Plant Development-I1 \ b) Vascular cambium consist two types of cell: Fusiform initials and ray Initials The fusiform Initials: are elongated cell with tapering ends. . They are found in tracheary elen~ents. fibres. vi) wood is agood material for carving idols. railway wagons. ix) cosmetics. The horizontal system is made up of xylem rays and the vertical consists of tracheary elements. cabinets. Usually itlooks like a convex lens both internally and externally. furniture. matches. . Ray Initials: are much smaller than fusiform initials and are aln~ost isodiameteric. c) In timber trade wood of dicotyledonous is known as hardwood and the wood as of gymnospern~s soft wood. viii) several music instruments. Woods have several uses which are listed below: building material for houses in windows. perfumes and incense sticks are made from oil derived from woods. The radial walls are thicker than the tangential wall.2. iv) electric poles. bridges. fibres and wood parenchyma. The living cells of rays are usually interconnected and a continuous system of living cells is formed. v) various aspects of art are also made up of wood. inlay work. the cell wall possess primary pit fields with plasmodesmata. soaps. masts. iii) bodies of auton~obiles. True (c). But these words do not express the degree of hardness.SAQ 5: a True b False c False. On stem they are longitudinally or horizontally arranged but most of the time they are scattered all over the entire surface. They have intense vacuolation. c) See section 10. Multi seriate: When it is more than two cell wide. True. railway sleepers. These woods are dense and heavy. xylem and phloem parenchyma and sieve elements. when the ray is one cell wide it is called uniseriate ray. SAQ 6: a (ii) b (iv) c(i) d(iii). But histologically the dicotyledonous woods has vessels while gymnospennous wood lack then. SAQ 3: a) . False (b). The large position of the wood is made up of fibers or fiber tracheids and they give strength to wood.
It will always remain at the same height above the ground. Dracaena. masses of complementary cells are pushed out. Abnormaly due to the formation of accessory cambium and its activity. ' 2. i Stem increases in height due to the act~vity shoot apical meristem. Examples: Families.SAQ 7: Lenticels: Restricted areas of relatively loosely arranged cells. SAQ 8: Cambial variants are of following types: i) ii) iii) iv) The cambial is persistent and normal in position but its products show unusual arrangement and proportion. In the dicotyledons the stem generally increases in girth due to the activity of the vascular cambium. As the division progresses. and you can see the differences between the activities of the apical meristem and of the vascular cambium. larger size of the numerous component cells. Lenticels usually found on young branches of stem and roots. Bougainvillea. Then there is formation of several concentric cambia. Examples: Most of the dicoty 3. Agave etc 2. They protude above the periderm because of the loose arrangement. Anomally due to the formation of interxylary phloem as a result of aberrant activity and position of cambium. Some of these parenchyma elements differentiates into vascular. Dicot Stem 1.The nail may eventually become embedded as the stem increases in girth. Bundles are also separated from one another by radial panels of parenchyma. Though the cambia are continuous they produce separate bundles which are surrounded by conjuctive tissue. Thus alternate bands of proliferated pericycle and vascular bundles are formed which is seen as dark coloured and light coloured rings respectively. The growth is known as secondary growth or secondary thickening. Secondary Growth SAQ 9: In the beet roots first formed cambium activity ceases very soon. All the cambial layers continue to function and produce a large amount of storage parenchyma and strands of xylem and phloem. bundles and remain embedded in the parenchyma or conjuctive tlssues.and rise above the surface of an organ. 3. 2. Monocot Stem _ . _ _ _ > Lateral meristem joins and makes vascular cambium which in turn cuts off secondary tissues. y ledonous woods plants such as Hetianthus. Complementary Cells: The cells which are derived from the divisions of the substomatal cells as well as those produced towards the exterior by the phellogen of the lenticels. They are involved in exchange of gases between internal tissues and the atmosphere through periderm.So if a of nail is driven in trunk. The cambium is abnormal in position but normal in activity. Answers to TQs: 1. The cambium cuts secondary xylem inside and secondary phloem outside. In some monocotyledons seems it is secondary meristem Vascular cambium produces only parenchyma. 1. . in the periderm.
A new phellogen differentiates a few millimeters within the cortex. resins. Tyloses totally block the cell.2. consisting of bundles embedded in nonvascular tissue. See section 10. gum. Interxylary phloem is found in following plants. See section 10. See section 10. This cambium divides rapidly and in about 10 years it forms sufficiently thick cork for commercial uses. The first formed periderm is removed when tree is about 20 years old and 40 cm in diameter. The bundles may be arranged in concentric rings or irregularly.8. and somewhat radially rounded. - 5. Some time unusual secondary growth occurs due to the formation of acessory cambia and their activity. Most of the commercial cork comes from Quercus fuber. 4. The cambium forms tissues centripetally. Heartwood is formed after disintegration of the protoplast. See section 10. and radially flatterned. There are two types of vascular tissues (i) a hollow cylinder or (ii) irregularly arranged bundles. Some cells are hollow and thinwalled. produced by the phellogen. 7. Phellogen abaxially cuts off derivatives as phellem. See Section 10. iii) The phelloderm-parenchymatous tissues in some species. Cork cells are dead.10. It is divided into three parts.This is known as heartwood. elasticity and lightness and see section 10. 6.5. The later type of cells may be often filled with dark resiniferous or tanninferous substance. In nearly all the trees the central portion consists of dead parenchyma which ceases to conduct water. hydrolysis of stored reserve materials and'formation of tyloses. 11.2. loss of cell sap. produced by the phellogen. 10 A vicennia Thunbergia Boilgainvillea Salvadora. tanins are accumulate in the heartwood making it extremely durable. Some others are thick walled and radially flattened. . Oils.9. In this species phellogen arises in the epidermis.6. i) The phellogen cork cambium ii) The phelloderm-parenchymatous tissues in some species.5. The phellem or cork cells are usually polygonal. The cells are arranged in compact radial rows which are devoid of intercellular spaces. Cork is of commercial value because it is impervious to gases and liquids and has strength. 9. Their cambial activity ceases and outside a bundle in the pericycle a new secondary cambium arises.8. Cork is an important part of secondary tissue which is termed as peridenn. Thus heartwood is preferred to sapwood for making furniture. The outer part of the secondary xylem with living parenchyma is named sapwood.Plant Development-ll 3. 8.