206 _Caravan’s Text Book of Botany - Paper B Chapter 16 COMPLEX TISSUES The complex tissues are heterogeneous in nature, i.e., these are compo: different types of cells performing diverse functions. Xylem and Phloem are the m: a complex tissues which together form Vascular Bundles of higher plants and are known ae as Vascular Tissues. XYLEM Itis a group of cells which are similar in origin and function but of more than one type in structure. The xylem is the principal water conducting tissue and along with the phloem t .e vascular tissue of higher plants. The xylem is comparatively more sclerefed constitutes th ides support to the plants; therefore, it and durable than that of phloem. The xylem also provi is member of the supporting tissue of the plant body s it consists of several types of living and non living cel ary Elements, i.e., the Tracheids and the Vessel Members which are the non-living cells that are principally concerned with transport of water and which also to a certain degree have 2 supporting function. Fibres are present in the xylem where they are mainly concerned with the strengthening of the plant body. Sometimes’ Sclereids may be present. Parenchyma Cells which perform storage and other functions also occur in the xylem. The xylem of some plants contains resin ducts. Xylem is a complex tissue a The most important cells are the Trache: Developmental, two types of xylem are recognized:- 4. Primary Xylem, derived from the procambium during the formation of pr plant body, developing from embryo. 2. Secondary Xylem, formed from the cambium, during second sta development, when increase in thickness (secondary growth) takes place: rimary ge of pat Elements of Xylem The following different elements are found in the xylem:- | 4. Tracheary Elements ied The term tracheary elements are derived from ‘trachea", a name originally PP! m elements resembling insect tracheae. certain primary xylet Two fundamental types of tracheary elements occur in xylem:(a) Trach (b) Vessel Members or Vessel Elements, @ tracheids: The tracheid is an elongate tube like cell having tapering, 1d lignified walls. The walls are not much thickened, nats erst and non-living at maturity. These are fundamental protop’2 plants. . In transverse section the tracheid is typically angular, though more or less rounded ; also occur. The tracheids of secondary xylem have fewer skies and are More sharply forms 1 Jar than the tracheids of the primary xylem. The end of a tracheid of secondary xylem is ul sanewat chisel-like. founded or oval ends and The tracheids are without cells of xylem of vascular =e ea oT ye fv tat ot | Member Fig. 17-2: A Vess*! A Fig. 17-4 A Tracheid & a i208 Caravan’s Text Book of Botany — Paper B The tracheids are dead empty cells and their walls have many borg arranged in rows. The lumen (cell cavity) of a tracheid is large and empty, ie, wit’ content. Out any Tracheids alone make the xylem of ferns and gymnosperms, while in the angiosperms they occur associated with the vessels and other xylary elements, The tracheids are specially adapted to the function of conduction. The thick ang walls of tracheids also aid in support and where there are no fibres or other support, the tracheids play prominent part in the support of an organ. 9 ele, em y (b) Vessel Members The vessels are long, cylindrical, tube-like with _lignified walls, and are foung mainly in the wood of angiosperms. They are formed by the absorption of end walls from 5 row of cells placed end to end, which are termed Vessel Members. Perforation Plates: The vessel members are usually perforated on the end wals | but sometimes the perforations arc formed on side walls also. Those parts of the cell wal ha bear perforations are called Perforation Plates. The perforation plates may contain one lage perforation and such perforation plate is termed as Simple Perforation Plate, or it may contin numerous perforations and is called Multiple Perforation Plate. The perforations may be arranged variously in a multiple perforation plate, e.g., perforations may be elongated and ae arranged in a parallel series, such plate is called a Scalariform Perforation Plate, or arrangst in a reticulate manner (forming a network), such plate is termed as Reticulate Perforaton Plate, or when the perforations are almost circular, the plate is known as a Foraminae Perforation Plate. Pteridium Fig. 17-3: Types of Perforation Plates. 4-Simple; 2-Scalariform; 3-Reticulate; 4-Forminate iSe SP SS ooo. unetion: Due to structural peculiarities i., presence ficient in transport of water and minerals 4 Of perforation plate, vessels chanical strength to the plant body, se al ate more prose ™ ‘Anatomy 209 ‘S compared to tracheids. These also Comparison of Tracheid and Vessel Tracheid 4, The tracheids are short and generally 1 mm. in length. In rare cases their length becomes upto 12.cm. - 2, It consists of a single elongated cell which possesses tapering end walls. 3, The tracheids are not tubular. Thése ate arranged in overlapping fashion and are separated by cross walls having Vessel 1. They are comparatively ly longer and mi reach upto 10 cm. in length. In rare aoe they attain the length upto 2-6 meters, e.g in Eucalyptus, Quercus. a 2. The vessel consists of a row of cells placed one above the other. Their intervening walls are absent 3. The vessels are tubular and have no cross walls. They may be perforated having many or a single pore. These-are bordered pits. adapted for conduction of water. 2. Fibres: The fibres associated with xylem are termed as Xylary Fibres. These fibres have thick walls and reduced pit borders as compared with tracheids from which they have evolved. Two main types of xylary fibres are recognised: |. Fibre-Tracheids and II. Libriform Fibres ther and also with the Fibre-tracheids and libriform fibres intergraded with each o! tracheids; therefore, the two kinds of elements are sometimes grouped together under the term "imperforate Tracheary Elements". These two main types of fibres can be distinguished on the basi and the amount and types of pits. is of wall thickness Fibre-Tracheids hose of libriform fibres, They have walls of medium thickness, i., not as thick a5 ¥ 16 ered pis. TY i {heker than those of tracheids. The fiber-tracheids have reduce Smaller pit chamber as compared to that of the tracheids. the secon’ rotoplasts 4 inal wall ; fate Fibre-Trachel is mature and separated by thin may qi Certain fibre-tracheid the protoplast persists afte? 2 Rien to Produce two or more protoplasts. These Pr tt Patttion walls and remain enclosed within the 2°! S ate called Gelatinous or Mucilaginous Fibres or Sept ~~ ‘Such fibres- and fibre- ids. The inner210__ Caravan’s Text Book of Botany ~ Paper B layer of secondary wall is rich in cellulose and poor in lignin i in these fit = absorbs water and may swells up to fill the entire lumen of the fibre bres. This Ly Libriform Fibres (liber— phloem fibres — similar to phloem fibres), They posses very thick walls with reduced simple pits. Libriform fibres are mostly met in woody dicots especially in more specialised families, e.g. in Leguminosae. Origin: Ontogenetically it is assumed that the fibres have developed from tracheids. Some intermediate forms found in some angiosperms, e.g., in Quercus spp., support this assumption. The steps towards these changes are- 1. Thickening of the wall resulting in decrease of lumen. 2 Reduction in number of pits and size of pitchambers leading to f | disappearance of bordered pits, 3. The shortening of cells. “However, in natural tissues of the plants, fibres are larger in size than those of tracheids. This happens ‘due to additional growth of the ends of the fibres and 7 sometimes due to fusion of fibres Function: 4. The xylary fibres —_ provide mechanical strength to specified parts of the plant body 2. Some of fibre elements are also concerned with storage. Fig. 17-5: A-Fiber-Tracheid; B-Libriform Fiber; C-Septate Fiber 3. Commercially, the fibres are used in textile arid cordage industry as brush fibres and filling fibres. 4, Parenchyma Cells: i imary ® Living parenchyma cells or wood parenchyma cells occur both in pri secondary xylem. These are called Xylem Parenchyma Cells.j In secondary xylem the parenchyma is found in two fc forms: i Axial Parenchyma: am tis derived from fusiform cambial initials alo ng with trachea ry elements and fibres. - be as long as the fusiform i These cells- may 7 O initials (fusiform pare: ‘shorter than the fusiform initials. If a tistorn dertvatve dines tenon les transversely many times pefore differ occur More more or less | (ii) Ray Parenchyma: The ray parenchyma Is formed by the ray initials of im. The ray parenchyma cells are arranged in the cambiul radial transverse series and form Xylem Rays. The ray parenchyma cells vary in shape. The axial parenchyma cells and the ray cells of the secondary xylem may or may not have secondary walls. If a secondary wall is present, the pit-pairs between the parenchyma cells and the tracheary elements may be bordered, half-bordered or simple. Only simple pit-pairs occur between parenchyma cells. Function 1. The xylem parent storage of food reserves In the form fat. ichyma cells are noted for of starch or reserves accumulate wth season and are al activity of the 2. Generally, the starch towards the end of the gro depleted during the cambi following season 3. Tannins, crystals and various other substances also occur in parenchyma cells nd ray Tyloses: In many plants, the axial a enter par Se ne eer that ese auilgron when these become injured or inactive. Vloses, ere from parenchyma cells are called Pairs connecting development occurs through the pit- elemente. "9 the parenchyma cells with tracheary Fig 47-7: A vesst entiation into parenchyma, it forms parench yma str commonly than parenchyma cells. The axial usar ae Svan) Jongated and placed end to end in the secondary xylem Is occur vertically el with Tyloses212 _Caravan’s Text Book of Botany — Paper B 10. 11. : Depending upon Comparison of Primary & Secondary Xylem Primary Xylem It is formed by the activity of procambium of the apical meristem. Protoxylem and metaxylem are distinct. position of protoxyim, it may be endarch, exarch or mesarch. The tracheids and vessels are relatively narrow and longer. Medullary rays found in this region are formed by the histogen of the apical meristem. Axial and radial systems arc; not distinct. Tyloses are not found in the vessels. There is no distinction into heart- ‘wood and sap-wood. Annual rings are not found. Medullary rays are always homocellular, i.e. each consists of only one type of cells. Xylem fibres are either absent and if present, these are, very few in number. PHLOEM The phloem is the principal food conducting tissue of the vascular plants. It is @ gor of cells which do not divide and are similar in origin and function but structurally of more ‘one types. The phloem and xylem are present together throughout the plant body. nature of the tisst The tissue was first recognised due to nature of its fibres which were used for and received the name of Bast. Harting in 1837 discovered sieve elements an ue was revealed. In 1858, Nageli gave it the name Phloem (derived fro! Greek word for bark). Secondary Xylem 1... It develops from vascular car which is a lateral meristem, oe 2. There is no distinction be protoxylem and metaxylem. = 3. Such distinction in xylem j is observed. ‘a 4. Both are normally wide and short 5. The secondary medullary arrays arise by the activity of ray initials of the cambial ring 6. The two are clearly distinct and well marked. 7. Vessels show tyloses. 8. Sap-wood and heart-wood are clearly demarcated in woody trees. 9. Annual rings are clearly distinct in the wood of temperate trees and shrubs. 10. Medullary rays may be homocelluiat or heterocellular. 11. Xylem fibres are found abundantly. binding the tm yn thefs me of appearance in le, the phloem is classified as p PMent of the as a whore “ ; iS Primar Plant or. 1 ge" bem is initiated in the embryo, is constantly added duinecondary Phloem, The 1 eon body, and completes its differentiation when the primar ‘evelopment of the win i gifferentiates from the procambium like af 16. Plant body is : xylem. The seconday Y fs fully pe cular cambium during Secondary growth nth in we fom loem differ rior ofthe stem or root | tion external to the xylem i i tly the phloem occupies @ posi xylem in the axis, or ab: Mosthe axis) in the leaves, but in certain tems and many dicotyledonous fantice (evay fro eae, Convolaceae, Solanaceae) a part of the phloem is located on the inner side mem or is adaxial (towards the axis). The phloe the * M present outside the xylem is called | Phloem or Abaxial Phloem. The phloem present inside the xylem is termed as eternal oem or Adaxial Phloem. Sometimes the phloem strands or layers are included in imal PRIOGn Myiem of certain dicotyledons, such phloem is termed as Interxytary or e secondary mF the included phloem appears in layers alternating with xylem layer, itis Induded ite ic and if it appears in strands surrounded by xylem tissue, it is known as caled SO aes the internal phloem is initiated somewhat later than the external phloem fai i not increased in amount by cambial activity. Elements of the Phloem Anatomy 213 elation to the develo, The phloem is a complex tissue comprising of more than one type of cells: The e pl i {ifferent elements found in the phloem are as follows: b> SIEVE ELEMENTS SIEVE TUBE PLASTIOS PARENCHYMA: PLASTID Sieve PLATE m Fia. 17-9 : Elements of Phloe! |244 _Caravan’s Text Book of Botany ~ Paper B 41. Sieve Elements: The conducting elements of the phloem are collective Sieve Elements and consist of: ()) less specialized Sieve Cells and (i) more gpg, tt Sieve-Tube Members or Sieve-Tube Elements. The sieve elements are thin waleg cells. The mature sieve elements lose their nuclei. The most characteristic feature of wey ve elements is the presence of Sieve Areas in their walls. _ i) Sieve Cells: These are narrow, elongated cells without prominent sieve ar The sieve calls taper at their ends-or have greatly inclined walls that overlap in the 4° They are found in lower vascular plants and gymnosperms. Sue, (ii) Sieve Tube-Members: These are long, tube-like, slender cells with Proming sieve areas (perforations like those of a sieve) in their end walls. These join end to eng form a vertical row of elongated cells called Sieve Tube. ALW1d 3A3IS GNNOdWOO Fig. 17-10 : 1-A Sieve Cell; 2-A Sieve Tube with Companion Cell Sieve Areas and Sieve Plates: af The sieve areas are considered to be modified firm pit fields and aera depressions in the walls in which groups of pores are located. Stra like protongatioAnatomy 215 Present in the sieve areas, Thi ese Connect the - The sieve areas can be distin 3-structure of Sieve Areas The sieve areas show variations in density and arrangement on sieve elements. Those parts of the cell wall that bear specialised sieve areas are called Sieve Plates, Simple Sieve Plate: If a sieve plate consists of a single sieve area, it is a Simple Sieve Plate (Cucurbita). Compound Sieve Plate: In certain elements, e.g., in Viti, the sieve plate contains i . itis ny sieve areas which may be arranged in scalariform. reticulate or any other manner. ma a Compound Sieve Plate. a A comers rmawentous renew seve nate SIEVE TUBE weMBER Fig. 17-11 : Ultra-structure of Sieve Plate e element and The Sieve Cell have unspecialized areas that are similar Tee seer Parts cannot be easily distinguished as sieve plates. The a Seer id slender, and they taper at their ends or have steeply in ee are towed ach other in the tissue and the sieve areas are more es on Se rien ular cryptogams and gymnosperms. Sieve-Tube Member: ee eee f the sieve areas are more highly specialised than ee tiger horionts © plates. The sieve plates are usually found on end walls t! ‘SVE tube members join end to end to form Sieve Tubes. the wat long an Overlap '" vase Some of F siey ie si216 Caravan's Text Book of Botany — Paper B Wall Structure & Protoplast: pectin. The thickness of the wall varies from species to species. In some species the homogeneous, while in others the wall is composed of two layers, a thin layer close middle lamella and a thicker layer next to the cytoplasm. The inner layer is lustrous The walls of sieve elements are usually primary and consist mainly of cellu lose ang Wall ig to the and ig termed as Nacreous Layer..In one group of conifers, true secondary wall has teen interpreted. Comparison of Vessels & Sieve Tubes Asieve tube differs from a vessel in certain respects, such as: Vessels Vessels have open ends Vessels are wider. These have thick, rigid and lignified walls, These have various thickenings on their walls. types of hysiological Comparison 5. 10. 12. They are dead when mature, but functional. These are permeable to both solute and solvent. They have low sap concentration. They do not have turgor pressure. They are partially collapsed when functioning. They absorb water or air when cut. They translocate both solutes and solvents. Their translocation speed is upto 75 em./min. 10. 11. Sieve tubes Sieve Plates are present at the ends of sieves tubes These are very narrow. These have thin, more extensible and cellulosic cell wall No such specialised thickenings ae found. They are alive when mature ard functional These are semi permeable in natue They have high sap concentration The turgid cells have high wo" pressure. They are distended by when — functioning. pressu® They exude cell sap when cut They translocate solute on! 5 is ue? Their translocation speed m/min (maximum). ‘oadin, 3. A of par and yy eleme te gl tissue Syste the paeae Anatomy 217, 2. Companion & Albuminous Cells: specialised types of parenchyma cells calle: Si sievertube members of angiosperms. The sala Sls afe associated wih the | Gntogenetically, a8 these develop from the same meristematic ell Such apis at | Shades longitudinally resulting in larger cell, that specialises into si Se Emersons coe | other develops into companion cell. The companion cells may eee rember and the bets to which they are related or they may be shorter, The compan ate ‘on various sides of the sieve tube or-they may form longitudinal iat ee ae sompanion cells are strongly attached’ to sieve-tube members and ane ny speed usually even by maceration. The walls between the companion cells ond ihe sieve ns members are thin and contain depressed areas, the primary pit fields Pesfhocemot ae seen in these walls under electron microscope. The companion cell retains its face maturity, is very rich in ribosomes, and contains numerous mitochondria, rough ER Ee pistids. Some companion cells contain P-Protein, The companion cells play an important partin the maintenance of a pressure gradient in the sieve tubes. In gymnosperms companion cells do not occur, but certain ray and phloem parenchyma cells are closely associated morphologically and physiologically with the sieve cells in conifers and Ginkgo. These cells are termed as Albuminous Cells. These cells stain deeply with cytoplasmic stains and are considered to be rich in protein: In secondary phloem uminous cells related to sieve cells ‘ontogenetically are found. The albuminous of Ephedra, albt tele can be distinguished from other parenchyma cells by the presence of connections with ieve cells are sieve cells ‘and absence of starch. The albuminous cells die. when the si disorganized. Highly respiratory and acid phosphatase activities occur in albuminous cells sesocated with sieve cells. The increased activity is restricted to those periods during which ieading and unloading of the sieve cells is taking place. 3. Phloem Parenchyma Cells: the phloem contains variable number ials and accumulation of tannins er cells as the sieve Ibuminous cells, torage of food materi arise from the same moth formed. The parenchyma cells of primary phloem lel with the longitudinal extent of the vascular stems, the Axial and Ray ™m Parenchyma whereas Apart from companion cells and al of parenchyma cells concerned with si and resins. The parenchyma cells may elements before the companion cells are are elongated and their long axes are para tissue. In secondary phloem, the parenchyma cells occur in two sys System. The parenchyma of the axial system is called ‘Axial Phioe! the parenchyma of the ray system constitutes Phloem Rays 4. Fibres: fae occur both in primary and seco a from fusiform eambial cells as components of the axial syste rete are usvall im are shorter than the fibres of primary phloem. The pits in ‘and act aS io Ple. In some plants the fibres of the secondary phloem act as con bres are thick walled "28 alter the sieve elements cease to-function. In some cases the and a ct as mechanical elements. ondary phloem, the fibres em. In se a vm. The fibres of secondary218 _Caravan’s Text Book of Botany - Paper B Primary Phloem The primary phloem develops from the procambium and may be divided ny Protophloem and Metaphloem. “ The Protophloem constitutes the conducting tissue of elongating plant parts ang contains sieve elements. In gymnosperm, the sieve areas are not distinguished in the protophioem elements. In angiosperms, sieve tube-members are found in protophloem of foots, stems and leaves while the companion cells are lacking. These sieve tube-memberg care long and narrow, and sieve areas can be distinguished only with difficulty. The sieve tube-members of protophloem remain active for a short period of time because (hey lack nucleus and cannot keep pace with the actively elongating plant organ and are passively stretched. The remnants of stretched sieve elements completely disappear later on and the phenomenon is called Obliteration. In many dicot stems, the parenchyma of the protophloem becomes fibres after the obliteration of sieve elements. In leaves they form elongated collenchyma cells. ENDODERMIS ‘PRIMARY PHLOEM Fig. 17-12: Primary Phloem The Metaphloem matures after the growth in length of the surrounding tissue s complete. Therefore, it is retained as conducting tissue longer than the protophioem. In Oe where secondary growth does not take place, the metaphloem tissue act aS cond tissue, whereas in plants with secondary growth the metaphloem sieve elements be inactive after the formation of secondary conducting elements. In these cee d metaphloem may be partly crushed or completely obliterated. The sieve elem metaphioem are longer and wider than those of protophloem, and their sieve areas O° itinet, In metaphioem of monocots. Sieve tube- members and companion cells af Pe while in metaphloem of dicots, the sieve-tube members, companion cells and PAnatomy 219 Parenchyma is found. ic phloem in di found. The metaphloem of dicots usually lack fibres. If fibres " ‘occur in prim: ots, these are in protophioem, ea ally distinct in erge gradually. In plants having secondary The arrangement of the secondary xylem. In Phloem, al: fusiform initials of the cam| the cambium, are present elements of secondai Iso, two systems, ie. ium and Horizontal or Ray ry phloem is parallel to that of the » the Vertical System derived from System derived from the ray initials of The principal components of Vertical System of phloem are sieve elements, phloem parenchyma and Phloem fibres. The Horizontal System comprises of ray parenchyma cells. Storied, non-storied and intermediate arrangements of the elements may be found in different ngement of the tissue is determined by the nature of the cambiym, ie., whether it is storied or Non-storied, and by the degree of elongation of various elements of axial system during tissue differentiation In many species of Woody dicots, Growth Rings are observed in phloem but they are less distinct than those of xylem. These growth rings are formed due to differences in cells produced at the beginning and end of the growth season. The cells formed at the beginning of the season extend radially while those produced at the end of the season are flattened. The growth rings become obscure due to obliteration of sieve elements and enlargement of parenchyma cells. In many gymnosperms and angiosperms tangential bands of fibres develop in secondary phloem. init lis towards both the xylem and the phloem, The cambial ray initials produce cel rs oF Spe a Fs pees 's and the phloem rays are continuous. Ease vestid ium, the xylem and phloem rays are constitute the Vascular Ray. In the vicinity of ee cambium, i a an oom ye in many plants the mature outer } ma wath ricea ae are uniseriate, biseriate or multiseriate; they vary in heat sd anal sn ‘large rays may be present in the same species. The phloem rays = ee ee rays as aie cambium produces less phloem than xylem and also f phellogen hl through the activity of ene Sousa of . ly simple. The vertical system comprises The secondary phloem of Conifers is relativel Jig and in many plants fibres are also . ‘ jinous cel ells. ot sieves cells, parenchyma cells’ and aveualy uriseiate and consist of parenchyma ct Present. The phloem rays of conifers are us! t their ends and each are in Only. The sieve cells are slender, elongated, overlap om one et wero =n viongtudrl stands end soe eaercealt times of the year Sr £18 occur in longitudinal strands and store starch al in eo yo nn Contain resins, fats and tannins. The secondary P &; rs of Pinaceae. “"als. Some species lack fibres, €.g., membe' bers jeve tube-mem! d phloem with elongated © ry phloem fibres of Many woody dicots have pore rand walls. THE seoon im sind include i in e ously arranned ASPaper B " scattered groups of other phloem elements, or may be arranged in tanga aiternate with bands of the sieve tubes, companion cells and phloem pares ig scattered among the rest of elements of the vertical system. The phloem fibres chyna from the parenchyma cells of the non-functioning phloem are called Pius deve, Sclereids occur both in functioning and non-functioning phloem where renee parenchyma cells. The arrangement of sieve-lube members and the parenchyma n° fron in various plants. These may form alternating separate bands or may be arranges ite, "ati rows. NON-FUNCTIONING PHLOEM FUNCTIONING PHLOEM: PART OF RAY SIEVE TUBES COMPANIONT LL st He J T a PART OF RAY FIBRES CRYSTALS ape EARLY SEASON PHLOEM Li Fig. 17.13: Secondary Phloem Comparison of Primary and Secondary Phloem o bi 1. It develops from procambium apical 1. It develops from vascular 63! meristem. which is a lateral meristem 2. There is distinction between first 2 Such distinction is not found, protophloem and later formed metaphloem. developedPhioem fibres are found in the outer part of phloem. sieve tubes are usually longer with narrow lumen. phioem parenchyma is poorly developed. Sclereids are usually not found. Medullary rays found in this region are developed from the activity of apical meristem. Medullay rays are homocellullar. Callus formation around sieve pore is either very little or even absent. Phloem fibres are ‘found: samon Phloem Parenchyma cells, Sieve tubes are -sho rer with broad lumen ° Phloem Parenchyma are abundant and are concerned with Storage of food materials or may contain crystals, latex etc. : Sclereids are found in secondary phloem of several plants. Secondary medullary rays found in this region arise by the activity of ray initials of carnbium. Medullary rays may be homoceltular or heterocellular. Callus formation is abundant.