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Plant Tissues Lesson Prepared Under MHRD project “National Mission on Education Through ICT” Discipline: Botany Paper: Plant Anatomy National Coordinator: Prof. S.C. Bhatla Lesson: Plant Tissues Lesson Developer: Dr. Arun Kumar Maurya and Dr. Anita Rani Department/College: Dayal Singh College Lesson Reviewer: Dr Basudha Sharma Department/College: MM (PG) College,Modinagar, Uttar Pradesh Language Editor: Dr Sonal Bhatnagar Department/College: Hindu College Lesson Editor: Dr Rama Sisodia, Fellow in Botany ILLL Institute of Life Long Learning, University of Delhi 0 Plant Tissues Table of Contents Chapter: Plant Tissues Introduction Classification of Tissues Simple tissue Parenchyma Distribution Shape and arrangement Structure and function Synthetic parenchyma Structural parenchyma Boundary parenchyma Transport parenchyma Medullary parenchyma Storage parenchyma Collenchyma Properties of collenchyma Types of collenchyma Functions of collenchyma Sclerenchyma ° ° Fibres Properties of fibres Distribution Type of fibres Function Sclereids Type of sclereids Function Institute of Life Long Learning, University of Delhi 1 Plant Tissues Complex Tissue/ Vascular Tissue ° ° Xylem Tracheids Vessel Xylem parenchyma Xylem fibres Phloem Sieve tube/Sieve cell Companion cell/albuminous cell/strasburger cell Phloem parenchyma Phloem fibres Summary Glossary Exercise Multiple choice questions References Institute of Life Long Learning, University of Delhi 2 Plant Tissues Introduction Cell is the smallest functional unit of life. In complex multicellular organisms, cells similar in structure and function are grouped together to form tissues. These tissues perform various functions of organs necessary for maintaining biological life. Thus, the plant body has a hierarchy of organs, tissues and cells. Classification of Tissues The wide variety in form and function of plant cells presents problems for the classification of cell types, tissues and tissue systems. Sometimes different types of cells have no well-defined boundaries between them and large numbers of intermediate forms can be seen. There is a continuum from one type of cells to other types. Some cells develop highly specialized, limited structures and functions, whereas others appear to carry out multiple functions and may even resume growth, cell division and differentiation. Some plant tissues are made up of only one type of cells called as “simple” tissues whereas others are composed of different cell types called as “complex” tissue. Thus, attempts have been made to classify tissues on the basis of mature structure, principal cell or tissue origin etc. Classification of plant cells and tissues are therefore artificial and should be viewed only for convenience of study and understanding. Institute of Life Long Learning, University of Delhi 3 Plant Tissues Figure: Diagrammatic representation of different types of plant tissues. Source: http://lurnq.com/lesson/Anatomy-of-Flowering-Plants-Part-I-Tissues/ Tissues are made up of multiple cells having a common origin and all of them together perform a similar or a set of similar functions. Depending on the capacity to divide, tissues are classified into main types: Meristematic tissues Permanent tissues Meristem is a group of actively dividing cells that do not form a specific organ but retained the capacity to divide and form new cells. Meristematic cells are compactly arranged without intercellular spaces. They are present at the apices of root and shoot (apical meristem), between xylem and phloem (lateral meristem) and at leaf and node bases (intercalary meristem) and do not store any reserve food material. All the permanent cells formed are product of meristematic tissue that form specific tissue or organ and are unable to divide after they attained a permanent shape and size. Plants are composed of three types of tissue system: dermal, vascular and ground/fundamental tissue. Each system is continuous throughout the plant body. (See next chapter) Ground tissue is mainly composed of three basic cell types: Parenchyma, Collenchyma and Sclerenchyma. Plant cells possess structural adaptations that make specific functions possible and hence differentiate them from each other. This differentiation is evident within the protoplast, cell contents and by modification of cell walls. Simple Tissues Parenchyma Parenchyma (Para means “beside” and chyma means “infilling”) is a versatile ground tissue that constitutes the “filler” tissue in soft parts of the plants. The term parenchyma was first introduced by Nehemiah Grew in 1682 (Metcalfe, 1979). Ontogenetically parenchyma is the precursor of other tissues. The body of the primitive organisms is always parenchymatous. These cells when compared morphologically, developmentally and physiologically with other complex tissues are found to be relatively undifferentiated and unspecialized. Institute of Life Long Learning, University of Delhi 4 Plant Tissues Figure: A) meristematic, B) parenchyma, C) collenchyma and D) sclerenchyma cells. Source: http://plantphys.info/plant_physiology/basiccytology1.shtml Distribution Parenchyma cells are present in almost all plant organs and forms the ground tissue in which other tissues are embedded. They are present in the cortex, pith of roots and stems, mesophyll of leaves, flesh of succulent roots, seed endosperm etc. Parenchyma cells are also present in epidermis and vascular tissues. Shape and arrangement Parenchyma cells are generally polyhedral i.e. with many sides. Even if the parenchyma cells are approximately isodiametric, they are not spherical but have many facets. Plant cells rarely approach this ideal from because inside the tissue the pressure exerted is uneven. Institute of Life Long Learning, University of Delhi 5 Plant Tissues Figure: Simple tissue: Parenchyma cells A) Transverse section B) Longitudinal section Source:http://lurnq.com/lesson/Anatomy-of-Flowering-Plants-Part-I-Tissues/ The parenchyma cells are closely packed in the endosperm of seeds where they are compactly arranged with no or few intercellular spaces. Storage parenchyma has abundant intercellular spaces in aquatic plants forming aerenchyma. Leaf mesophyll cells show extensive developed intercellular spaces, as a result of continued increase of cell volume during growth, the number of walls faces increases above fourteen. This makes it impossible for all the sides to remain in contact with all the sides of the neighboring cells thus, the intercellular space develops. It develops in two ways: By the separation of neighbouring cell walls, as in development of resin ducts in Pinus, known as schizogenous cavity. By the disintegration of the cells in the place where the space develops. Such development is known as lysigenous development. It is seen in the formation of oil cavities in the peel of citrus fruits. In some cases spaces are formed by involving both the methods and the development is called as Schizo-lysigenous development. The development of intercellular spaces in protoxylem is formed by such method. Institute of Life Long Learning, University of Delhi 6 Plant Tissues Figure: A) Pinus needle showing schizogenous development of resin duct B) Peel of Citrus fruit showing lysigenous development of oil gland or cavity. Source:http://www.biologie.uni-hamburg.de/bonline/library/webb/BOT410/Secretion/PinuSecEpithelium400.jpg;B) http://www.sbs.utexas.edu/mauseth/weblab/webchap9secretory/9.3-7.htm Parenchyma cells may be elongated columnar in shape as found in palisade tissue of the leaf. Stems of plants shows well-developed air spaces (Scirpus, petiole of Canna leaf) with their stretched arms, such parenchymatous cells are called stellate parenchyma. Parenchyma cells can be variously lobed e.g. spongy mesophyll and palisade parenchyma of Helium and Pinus needle. Parenchyma cells may have folded projections e.g. mesophyll of Xanthorrhoeaceae. Some parenchyma cells shows inner walls protuberance for short distance transfer of solutes called as transfer cell such as nectaries, salt gland etc. These protuberances increase the surface area for absorption or secretion. Such cells are called as transfer cells. Structure and function of parenchymatous cells It is least specialised plant cell. Thin and somewhat flexible primary cell walls. It is living at maturity and has a large central vacuole. It carries most of the plant metabolic function. Most parenchyma cells have the ability to resume meristematic activity and differentiate into other cell types under special conditions like during repair and Institute of Life Long Learning, University of Delhi 7 Plant Tissues replacement of organs after injury, regeneration, formation of adventitious roots and shoots union of grafts. Lobed parenchyma Figure: A) Transverse section of leaf of cattail (Typha) leaf showing stellate parenchyma. B) T.S. of needle leaf of pine (Pinus) showing presence of lobed parenchyma cells (all the red-stained cells). Source:A) http://www.sbs.utexas.edu/mauseth/weblab/webchap3par/3.3-6.htm B) http://www.sbs.utexas.edu/mauseth/weblab/webchap9secretory/9.3-6.htm Parenchyma cells usually have thin primary walls, but thick primary walls may also exist in some plants e.g. storage parenchyma cells of endosperm of Phoenix, Diospyros, Coffea and Asparagus, have very thick walls with hemicellulose, getting accumulated there as reserve substance. The walls of these cells gradually become thinner during germination as the reserve substance is used by the developing embryo. Parenchyma cells may also be induced to undergo lignification when infected by microorganisms suggesting the role in disease resistance. Parenchyma tissue is involved in almost all the physiological function of plant such as storage of food, provide turgidity to the softer parts and helps in slow conduction of various substances. They are known to perform specialized functions for which these cells undergo various modifications. As parenchyma cells shows various functions and structural modification these can be classified into various types:- Institute of Life Long Learning, University of Delhi 8 Plant Tissues Synthetic parenchyma Synthetic parenchyma cells are that synthesizes something: Photosynthetic: These are chlorophyll containing cells so called as chlorenchyma. It is found in the mesophyll leaves, some young stems, succulent stem, primitive plants, herbaceous plant etc. Mesophyll tissue is differentiated into compactly arranged columnar cells called as palisade and loosely arranged tissue called spongy parenchyma. Chlorenchyma cells are characterised by conspicuous vacuole and the cells have air spaces between them (lacunate) e.g. T.S. Nerium leaf. A B Figure: A) T.S. of Nerium leaf showing photosynthetic parenchyma cells showing both palisade and spongy parenchyma; B) Meristematic cells are parenchyma cells that are composed of immature cells with no intercellular spaces with abundant cytoplasm and one or more nuclei. Source:http://botit.botany.wisc.edu/Resources/Botany/Shoot/Leaf/Syringa/Cross%20Sec tion/Palisade%20parenchyma%20MC.jpg.html;B) http://commons.wikimedia.org/wiki/File:Mitosis_(261_13)_Pressed;_root_meristem_of_o nion_(cells_in_prophase,_metaphase,_anaphase,_telophase).jpg Meristamatic: These are immature cells contain abundant cytoplasm with one or more nuclei. They are parenchyma cells that consist of a group of cells which remain in continuous state of division or retain their power of division. They do not have intercellular spaces. The vacuoles are small or absent. These cells may differentiate into collenchyma and sclerenchyma or may stay as parenchyma. Institute of Life Long Learning, University of Delhi 9 Plant Tissues Secretory: Such parenchyma cells secrete substances and have dense protoplasts. They may contain plenty of ribosomes; golgi bodies or ER depending upon their secretion. Structural parenchyma This includes ground tissue, pith and cortex which is non-photosynthetic in nature. Figure: T.S. stem of Medicago showing ground tissue. Source:http://botit.botany.wisc.edu/Resources/Botany/Cells%20and%20Tissues/Medicag o/Labelled%20Medicago.jpg.html Aerenchyma: These are specialized parenchyma cells with large air spaces and cavities, such parenchyma tissue is known as aerenchyma. The aquatic plants provides buoyancy because of the aerenchyma in the cortex and through this the plant can float on the surface of water and also gaseous exchange takes place for e.g. leaf of Nymphaea, Hydrilla and Myriophyllum. Ethylene accumulates in waterlogged tissue and this gas induces programmed cell death and formation of aerenchyma. It may also be formed constitutively with no environmental stimulus e.g. rice roots. Institute of Life Long Learning, University of Delhi 10 Plant Tissues Figure: A) T.S. of Nymphaea leaf showing aerenchyma tissue; B) T.S. stem of Myriophyllum showing aerenchyma cells in the cortex region; C) T.S. of adventitious roots of (a) rice and (b) maize taken 50 mm from the root apex and showing lysigenous aerenchyma formation. Note the cubic cell packing in the rice cortex contrasting with the hexagonal packing in maize (Micrographs courtesy E. Armstrong). Source:A)http://student.nu.ac.th/cherrycoke/lesson6.htm; B)://www.uri.edu/cels/bio/plant_anatomy/26.html C)http://plantsinaction.science.uq.edu.au/edition1/?q=content/18-1-2-adaptiveresponses-waterlogging Institute of Life Long Learning, University of Delhi 11 Plant Tissues Boundary parenchyma Epidermis: It is the outer most layer in plant. Outer surface of epidermal cells has a layer called cuticle which is made up of waxy substance called cutin, due to this cuticular layer rate of transpiration is reduced. Figure: A) T.S. of leaf (a portion enlarged) showing epidermal cell covered by thick cuticle. B) Diagrammatic representation of endodermal cells bearing casparian strips which prevents the entry of substances into the vascular bundles. Source:http://www.studyblue.com/notes/note/n/midterm-1-definitions-withpics/deck/6021799; B) http://www.studyblue.com/notes/note/n/midterm-1-definitionswith-pics/deck/6021799 Endodermis: Inner most layer of the cortex in stem and roots is called as endodermis. It has a casparian strip around each cell which is made up of suberin. Endodermal cell prevents the entry of substances into the vascular cylinder. Figure: A) T. S. of Psilotum nudum rhizome haplostele (triangles indicate position of endodermis) B) T. S. root of Smilax showing exodermis. Institute of Life Long Learning, University of Delhi 12 Plant Tissues Source:A)http://www.phytoimages.siu.edu/imgs/Cusman1/r/Psilotaceae_Psilotum_nudum _41759.html; B) http://phytoimages.siu.edu/imgs/Cusman1/r/Smilacaceae_Smilax_sp_46545.html Exodermis: The cells of exodermis is similar to endodermis but one or more cell in depth in some roots, a type of hypodermis, the walls of exodermis may be suberized and /or lignified for e.g. In Smilax it is sclerenchymatous. Transport parenchyma The parenchymatous cells associated with xylem or phloem is meant for the transportation of water, minerals and food particles are known as transport parenchyma. Figure: T. S. stem of Cucurbita pepo showing primary xylem and phloem (the arrow indicates a sieve plate). Source:http://www.phytoimages.siu.edu/imgs/Cusman1/r/Cucurbitaceae_Cucurbita_pep o_45623.html Transfer cells with wall ingrowths called a labyrinth. In a variety of tissues where transport of solutes over short distances i.e. active transport or facilitated transport is required, surface area including cell membrane increases by wall ingrowths e.g. haustorial cells in parasitic plants such as Cuscuta. Institute of Life Long Learning, University of Delhi 13 Plant Tissues Figure: Cuscuta campestris: haustorium penetrating host tissues. Source:http://www.phytoimages.siu.edu/imgs/Cusman1/r/ConvolvulaceaeCuscuta_camp estris_47184.html Medullary parenchyma It is found radially arranged in between the vascular bundles in the stem and is meant for storage of reserve food. Storage parenchyma It occurs in many plants parts such as tubers, fruits etc. The parenchymatous cells of plastids which store starch is called amyloplasts. In petals of flowers the plastids become chromoplasts. Reserve materials are stored in parenchyma cells, i.e. in form of fluid in the vacuoles (e.g. amides and proteins) or in the form of small solid particles (starch, proteins, oils, fats etc.) or liquid in the cytoplasm. Cell sap presents in the roots cells of sugar beet and bulb scales of Allium cepa store amides, proteins and sugars. Cotyledonary cells of many species of legumes show proteins and starch grains in the cytoplasm. Parenchyma cells may be involved in the storage of water as in succulent plants. Such cells are usually large, thin walled with a thin layer of cytoplasm. The vacuole has somewhat mucilaginous sap which increases water holding capacity of the cell. Institute of Life Long Learning, University of Delhi 14 Plant Tissues Tannins may also be present in parenchyma cells, such cells may be scattered in the plant or may form continuous system. Tannin are found in vacuole. Parenchyma cells may store mineral substances in the form of crystals. Figure: Parenchyma cell showing A) chloroplast and B) chromoplasts C) starch grains (bean cotyledons) Source: http://student.nu.ac.th/cherrycoke/lesson6.htm B) http://www.studyblue.com/notes/note/n/midterm-1-definitions-withpics/deck/6021799 C) http://www.uri.edu/cels/bio/plant_anatomy/27.html Collenchyma Collenchyma (Greek word “Colla” means “glue”) is another group of simple tissue consisting of living cells found in plants. The first use of "collenchyma" was by Link (1837) who described it as the sticky substance found in the pollens of plants Bletia (Orchidaceae). Later on the term collenchyma was coined by Schleiden (1839). Collenchyma cells are elongated, soft pliable cells with non-lignified thickened primary walls. Sometime collenchyma cells contain numerous chloroplasts. Due to the presence of thick wall it has specialized to function as supporting tissue. Collenchyma cells retain active protoplast capable of removing the wall thickening when the cells are induced to resume meristematic activity. Institute of Life Long Learning, University of Delhi 15 Plant Tissues Properties of collenchyma Cell wall of collenchyma in addition to cellulose contains large amounts of pectin and hemicelluloses without lignin deposition. Presence of pectin makes them hydrophilic and helps in retaining much water. Ultrastructural detail shows presence of cross poly lamellate or helicoidal structure in collenchyma wall with primary pit fields. Collenchyma cells are known to possess several pattern in their wall thickening which can be seen either in the corners of the cell, inner and outer tangential wall or on two opposite wall. A B Figure: collenchyma cell A) transverse section B) longitudinal section. Source:http://lurnq.com/lesson/Anatomy-of-Flowering-Plants-Part-I-Tissues/ Types of collenchyma On the basis of thickening on tangential wall, three types of collenchyma cells are found such as plate or lamellar, angular and lacunar/lacunate collenchyma. Figure: Types of collenchyma cells. Institute of Life Long Learning, University of Delhi 16 Plant Tissues Source: http://student.nu.ac.th/cherrycoke/lesson6.htm Lamellar or Plate collenchyma When the wall thickening is mostly restricted to the tangential wall called as lamellar/ plate collenchyma. They are found in stem cortex of Sambucus nigra, Sanguisorba, Rheum and Eupatorium and in the petiole of Cochlearia armoracia. Figure: A) T.S of Sambucus stem showing Lamellar collenchyma B) Angular collenchyma in petiole of Cannabis Source: A) http://www.uri.edu/cels/bio/plant_anatomy/34.html B)http://www.doctortee.com/dsu/tiftickjian/cseimg/botany/plant-anat/stem/cannabisstem-xs.jpg Angular Collenchyma Angular collenchyma cells are those in which the wall thickening is restricted to the corners of cells and can be seen in the stem of Atropa belladonna, Solanum tuberosum and petiole of Begonia, Beta, Coleus, Cucurbita, Morus, Ricinus Vitis, Cannabis and Celery (Apium graveolens). Institute of Life Long Learning, University of Delhi 17 Plant Tissues Figure: Angular collenchyma in Celery petiole Source: http:// www.doctortee.com/dsu/tiftickjian/plantanat/collenchyma Lacunar Collenchyma In above two types intercellular spaces are absent but some time space gets developed in the angular type of collenchyma that give rise to special type of collenchyma known as lacunar / lacunate collenchyma e.g. are seen in adventitious root of corn, stem cortex of Brunellia and Salvia and also present in members of Asteraceae and Malvaceaae. Figure: A) Lacunar collenchyma from adventitious root of corn B) Annular collenchyma located below the central vascular bundle of the leaf Source: A) https://www.etsy.com/listing/80113232/pink-corn-root-8-x-10; B) http://www.ujaen.es/investiga/atlas/atlas_ingles/hojaolivo/hojaolivo100x3colenquima.ht m Annular collenchyma The annular collenchyma is characterized by cell wall that is more uniformly thickened and lumen is more or less circular in outline. It can be distinguished from angular collenchyma on the basis of degree of wall thickening restriction to the corners of cell. Angular collenchyma shows thickening only at the corner but in annular, thickening becomes massive and identity at corner thickenings becomes indistinguishable. Thus, the lumen assumes a circular outline in contrast to angular type collenchyma. Collenchyma cells show simultaneous increase in thickening and surface area. The cell wall may become modified in the older plant or can modified into sclerenchymatous cells by deposition of lignified secondary wall bearing simple pits. Institute of Life Long Learning, University of Delhi 18 Plant Tissues Functions of collenchyma The collenchymas provide flexibility, elasticity, mechanical support and strength to the tissue. It also plays an important role in photosynthetic activities due to the presence of chloroplasts. Sclerenchyma Sclerenchyma is a specialized tissue consisting of a group of cells in which secondary walls are often lignified. The term Sclerenchyma is derived from the Greek word ‘skleros’ means ‘hard’ and ‘enchyma’, an ‘infusion’. The term sclerenchyma was coined by Mettenius in 1805 and the cells are known as sclerenchymatous cells. Sclerenchyma cells may or may not retain their protoplast at maturity. On the basis of length, sclerenchyma cells are of two types; fibres and sclereids. Usually the fibres are long while sclereids are short sclerenchyma cells. Fibres Fibres are long, spindle shaped cells with thick secondary wall and occur as strands in plant. They exist as overlapping structure and impart strength to the fibre bundles. Rutting is the process for fibres extraction from plant body which separates the fibre bundles from associated non-fibrous cells. It is carried out in pond and this process is hastened and assisted by microorganisms. Properties of fibres The fibre cells have presence of high lignin content and absence of pectin and cellulose, their walls are not much hydrated or they have very less affinity for water. This property makes wall more elastic in nature than plastic. Fibres possess simple pits whereas border pits are relatively scarce. Distribution Fibres are distributed in plants as separate strands either in cortex and in phloem or as sheath or bundle caps associated with the vascular bundles or may be grouped or scattered in xylem and phloem. Monocot and dicot fibres show several characteristic patterns. In Poaceae, fibres form a system having the shapes of a ribbed hollow cylinder Institute of Life Long Learning, University of Delhi 19 Plant Tissues with the ribs connected to the epidermis. Vascular bundle have prominent sheaths of fibres and the peripheral bundles may be irregularly fused with each other or united by sclerefied parenchyma into a sclerenchymatous cylinder for e.g. Zea mays, Saccharaum, Andropogon, Sorghum. Figure: Showing location of different types of fibres Source: http://student.nu.ac.th/cherrycoke/lesson6.htm Type of fibres In angiosperms fibres are located in stem as outermost part of the primary phloem as an astomosing strands or tangential plates. On the basis of their position in plants they have been kept in two groups, xylary (intraxylary) and extraxylary fibres. The fibres present in xylem are called xylary or intraxylary fibres. Xylary fibres are also known as wood fibre and are of following types: a) Libriform fibres b) Fibre tracheids c) Septate fibres d) Mucilage fibres These libriform and fibre tracheids are classified on the basis of type of pits present on their walls. Libriform fibre (Latin Liber means inner bark) is longer and possesses simple pits whereas fibre-tracheids are shorter with bordered pits. Institute of Life Long Learning, University of Delhi 20 Plant Tissues Figure: Quercus maceration showing fibre-tracheids and libriform fibres Source: http://images.botany.org/set-17/17-104h.jpg Fibre cells generally loses their protoplasm and become dead at maturity but in many woody plants fibre retain their protoplasm and act as storage cell for carbohydrate and convert them into sugars when plant requires. Otherwise, their prime function is to provide mechanical support. Septa or cross wall formation takes place in phloem or xylem fibre of dicot species that undergoes regular mitotic division after secondary wall is deposited which leads to partitioning of fibre into two or more compartment. Such fibres are known as septate fibre. These fibres not only occur in dicot but also in some monocots such as Palmae and Bambusoideae. These fibres are non-vascular in origin. The septa include a middle lamellae and two primary wall that may or may not be lignified and remain in contact. The septate fibres of bamboos are characterized by thick polylamellate secondary walls with additional secondary wall lamellae. Septa of the fibre containing protoplast are interconnected by plasmodesmata, thus indicating role of septa in support and in addition to it they perform the storage function because they contain starch grains and sometimes crystals of calcium oxalate. Sometime fibre also contains gelatinous layer (G layer) which makes the innermost secondary wall layers, it contains high cellulose content but lignin is absent that distinguishes it from the outer secondary wall layers. Presence of cellulose makes the G layer hygroscopic and thus it swells up by absorbing large amount of water, it may occlude the lumen of the cell and upon drying it pulls away from the rest of the wall. They are also called as reaction fibre or mucilage fibre. These fibres are neither strictly xylary nor extarxylary as they have been found in xylem and phloem of roots, stem and leaves of dicots and in nonvascular tissue of monocot leaves. The role of G layers in Institute of Life Long Learning, University of Delhi 21 Plant Tissues leaves may be to assist in the maintaining the orientation of leaves with respects to gravity and display of the leaflet to the sun. Extarxylary fibres are located outside the xylem and these are of three types: a) Phloem fibres b) Pericyclic or perivascular fibre c) Cortical fibre Phloem fibres Phloem fibre also known as bast fibre which originates in early part of primary phloem but functions as fibres after their primary function i.e. conduction ceases and thus called as primary phloem fibre or protophloem fibre. Other examples of primary phloem fibre are stem of Sambucus (elderberry), Tillia (Basswood), Liriodendron (Tulip tree), Vitis (Grape vine), and Robinia pseudoacacia (Black locust). Flax fibres (Linum usitatissimum) exist as single band with several layers in depth and are located on the outer periphery of vascular bundle. When fibres are located within the secondary phloem called as secondary phloem fibres. Soft fibres are obtained from phloem fibre of eudicots and represent the bast fibre of commerce. They are soft, flexible and may or may not be lignified. The example of bast fibres are hemp (Cannabis sativa) used in cordage, jute (Linum usitatissimum) and remie (Boehmeria nivea) used in textile. Pericyclic or Perivascular fibre Perivascular fibres are extraxylary fibre found in stems of dicots, located in the periphery of vascular bundles inside the innermost cortical layer as in Aristolochia and Cucurbita. Extraxylary fibres also include the fibre of the monocot whether or not associated with the vascular bundles. They often have thick cell wall and variability seen in lignin deposition on cell walls. Cortical fibres are extraxylary fibre found in stem and originate in cortex e.g. Barley. Cortical fibre gives mechanical strengths to the plant body. The fibres obtained from monocots are basically obtained from leaves and are hard and stiff in nature, and thus they are called as hard or leaf fibre. In contrast to soft fibre, hard fibres are rich in lignin present on walls, for e.g. abaca or manila hemp (Musa textilis), bowstring hemp (Sansevieria sp.) and newzeland hemp (Phormium tenax). All of them are used in cordage making. Henequen and Sissal (Agave sp.) is used in cordage and Institute of Life Long Learning, University of Delhi 22 Plant Tissues coarse textile. Pineapple fibre (Ananas comosus) is used in textile. The plants like maize (Zea mays), sugarcane (Saccharum officinarum) and esparto grass (Stipa tenacissima) are used for making paper whereas leaf fibres in the xylem as raw material. Figure: The broken ring of perivascular (extraxylary) fibres in Aristolochia. Source:http://www.biologie.unihamburg.de/bonline/library/webb/BOT410/410Labs/Labs HTML-99/Stems-2/Labstm2-99.html Some cells in plants are not fibre but they act as fibre and thus are included in fibre such as cotton fibre which are obtained from the epidermal hairs of the seed of Gossypium. Some plant structure can also be used as fibre such as stem of rattan (Calamus palm) and Raffia is composed of leaf segments of raffia palm. Sclereids The term sclereid was coined by Tschierch in 1885. They are short sclerenchyma cells having thick and strongly lignified secondary walls with many simple pits. The secondary walls are multilayered. Some sclereids have thin secondary wall and contain living protoplast at maturity. Sclereid protects soft plant tissue from herbivores or mechanical damage. Institute of Life Long Learning, University of Delhi 23 Plant Tissues Type of Sclereids: Depending upon the basis of size and shape of sclereids, they have been classified into five main groups, a) Brachysclereids or stone cells b) Macrosclereids c) Osteosclereids d) Astrosclereids e) Trichosclereids Brachysclereids Brachysclereids are also known as stone cells. These are isodiametric or elongated cells and are distributed widely in cells of cortex, phloem and pith of stem and are also found in the flesh of food. They are present in fruits of Prunus, quinace (Cydonia); elongated sclereids are present in the endocarp region of apple seeds and stone fruits (Drupe); exocarp region of fruits of Manilkara achras; pulp portion of Mimusops elangi and Psidum guajava, cortex region of Cinnamomum zeylanicum and exocarp region of Moringa olefera. Macrosclereids Macrosclereids are elongated and rod like sclerenchyma cells which form palisade like epidermal layer in seed coats of legumes. They are well developed in exocarp region of Malus sylvestris. Osteosclereids Osteosclereids are columnar in shape but their ends become enlarge in such a way that it appears bone like structure. They are well distributed in the sub-epidermal layer of seed coat of some plants e.g. adaxial leaf surface of Nymphaea nouchali, Phillyrea latifolia, Hakia and Osmenthus contain osteosclereids. Institute of Life Long Learning, University of Delhi 24 Plant Tissues Figure: A) Fruit of Prunus showing Brachysclereids B) magnified stone cell (sclereid) from the juicy mesocarp of a 'Bartlett' pear C) Maceration of bean seed coat showing macrosclereids D) W.M. of Osteosclereids E) T.S. leaf of Nymphaea showing astrosclereids F) Banana leaf clearing showing trichosclereids. Source: A) http://lurnq.com/lesson/Anatomy-of-Flowering-Plants-Part-I-Tissues/ B) http://waynesword.palomar.edu/ecoph17.htm C) http://www.uri.edu/cels/bio/plant_anatomy/39.html Institute of Life Long Learning, University of Delhi 25 Plant Tissues D) http://www.biologie.unihamburg.de/bonline/library/webb/BOT410/anatweb/images /ParColSclr/MacroSclerEtc.jpg E) http://www.biologie.uni-hamburg.de/bonline/library/webb/BOT410/anatweb/images/ParColSclr/NymphLfXSLive2Scler.jpg F) http://botweb.uwsp.edu/anatomy/images/schlerenchyma/pages/Anat0040.htm Astrosclereids When the central body of cell develops arms or lobes like extension giving appearance of star are known as astrosclereids. They are found in leaf of eudicot, adaxial surface of leaf of N. cristata. Trichosclereids Tricosclereids are thin walled sclereids resembling hairs with branches. They are found on the adaxial surface of leaves of Olea europea and Banana leaf. Besides this two additional types of sclereids are also reported in plants: Filiform sclereids are long cylindrical cells similar to fibres and are found in palisade and spongy parenchyma of olive (Olea europaea) leaf. Figure: Filiform sclereid in leaf of Olea europaea. Source: http://images.botany.org/set-17/17-060h.jpg Fibre sclereids are the fibre that differentiates in phloem and have characteristic of both fibre and sclereids and thus named as fibre sclereids. They have been reported in secondary phloem of root and shoot of the rosette leaves of Arabidopsis thaliana. Sclereids are distributed in almost every organ of plant body ranging from epidermis, Institute of Life Long Learning, University of Delhi 26 Plant Tissues ground tissue to vascular tissue and occur singly or in cluster. When they occur as singly they are known as idioblast. Dicot leaves are rich in variety of sclereids but are absent in monocots. Two pattern of distribution of sclereids are mainly observed in dicots; terminal pattern and diffuse pattern. In terminal pattern they are confined to ends of small veins as seen in Hakea, Mouriria, Boronia and Arthrocnemum whereas, in diffuse pattern either they occur either solitary or in groups dispersed throughout the tissue without any spatial relationship to the vein endings. The examples of diffuse pattern are seen in Olea, Osmanthus, Pseudotsuga and Trochodendron. Foliar structure as found in clove scale of garlic (Allium sativum) the sclereids forms part of the entire epidermis. The plant species having well developed intercellular spaces or air chambers possess trichosclereids such as Monstera deliciosa, Nymphaea (water lily) and Nymphaea (Yellow pond lily). Seeds contain seed coat which is hard and this hardness is due to development of secondary wall in the epidermis and in the layers or layers beneath the epidermis e.g. seeds of bean (Phaseolus), pea (Pisum), soybean (Glycine max) contain columnar macrosclereids in epidermis and osteosclereides beneath the epidermis. The seed coat of coconut (Cocos nucifera) contains ramiform pitted sclereids. Figure: Stone cells or sclereids Source: http://student.nu.ac.th/cherrycoke/lesson6.htm Institute of Life Long Learning, University of Delhi 27 Plant Tissues Functions Sclereids provide mechanical strengths and plays important role in guiding light within mesophyll. They are responsible for gritty texture found in some fruits like pear. Complex tissue/ Vascular Tissue: A tissue that consists of several kinds of cells which functions together is called complex tissue. Xylem, phloem and vascular cambium (absent in monocot and lower plants) are example of complex tissue as they are involved in the water-conduction, transport of solutes and food material and are known as vascular tissue. The vascular plants also referred to as tracheophytes, which include seedless vascular plants of Lycopodiophyta (horsetails), Pteridophyta (ferns), gymnosperms and angiosperms. The terms tracheophyte (vascular plants) is given because of to the characteristic conducting element present in the xylem tissues known as the tracheary elements. Figure: Components in open and closed type of vascular bundles. Source: http://bioict.exteen.com/ Institute of Life Long Learning, University of Delhi 28 Plant Tissues Figure: A) Detailed structure of closed type vascular bundle B) Vascular bundle from Ranunculus stem (open type) Source:A)http://botit.botany.wisc.edu/Resources/Botany/Shoot/Stem/Zea%20stem/Vasc ular%20Bundle%20MC%20.jpg.html B)http://www.uri.edu/cels/bio/plant_anatomy/106.html Xylem Xylem (Greek word ‘xylos’= wood) is an example of complex tissue forming a part of vascular tissue. The term xylem was introduced by Nägeli in 1858. Xylem is mainly responsible for the conduction of water and mineral salts from roots to rest of the plant. Two type of xylem tissue have been demarcated in plants - primary xylem and secondary xylem. If the origin of xylem tissue has occurred from procambium of apical meristem, it is called as primary xylem and if it has occurred from vascular cambium the xylem is called as secondary xylem. The primary xylem develops early in the life of plant than secondary xylem. Presence and absence of cambium in vascular bundle develops open and closed type of Vascular bundles respectively. Xylem tissue consists of four kinds of cells – a) Tracheids b) Vessels or tracheae c) Xylem fibres d) Xylem parenchyma Institute of Life Long Learning, University of Delhi 29 Plant Tissues The term tracheid is derived from “trachea” because of its resemblance with insect tracheae. There are two types of tracheary elements in xylem, tracheids and vessel elements. Both are nonliving at maturity and more or less elongated cells. They have lignified secondary walls. The primary xylem develops earlier and are first formed elements called as protoxylem (from the Greek proto, first) and a later formed part are called as metaxylem (from the Greek meta, after or beyond). The protoxylem differentiates in parts of primary plant body where growth and differentiation has not yet completed. The protoxylem usually contains few tracheary elements (tracheids or vessel). The metaxylem begins to differentiate in the growing primary plant body and gets mature after elongation is completed and it contains more tracheary elements. Figure: Ranunculus root vascular cylinder showing the metaxylem (central last maturing xylem) with fully formed lignified secondary cell walls. Source:http://www.lima.ohiostate.edu/biology/archive/roots.html The secondary xylem is formed by a complex meristem known as vascular cambium, consisting of vertically elongated fusiform initials and squarish or horizontally (radially) elongated ray initials and such development distinguishes it from the architecture of primary xylem. The composition of secondary xylem found is more complex than the primary xylem in having a wider variety of cells in the angiosperms. Tracheids Tracheids are elongated cell with blunt ends, present along the long axis of the plant system. Phylogenetically the tracheids are most primitive type of cell found in xylem. Institute of Life Long Learning, University of Delhi 30 Plant Tissues These are imperforate or non-perforate cells having only pit-pairs on their common walls with lignified secondary wall. Tracheids are imperforate cells with bordered pits on their end walls. The tracheids contain simple pits along with bordered pits. They are arranged one above the other and are present in both primary and secondary xylem with broader lumen than that of fibres. This broader lumen in tracheids and occurrence of the pits helps in water conduction with dissolved mineral salts by acting as conduits in vascular plants. The presence of thick hard and lignified wall offer mechanical support to the plants. Sometimes an intermediate type of cell element is also found in vascular system known as fibre-tracheids (for detail consult fibre topic). Figure: Components of xylem (vessel, tracheids and fibre with secondary thickening) Source:http://elte.prompt.hu/sites/default/files/tananyagok/plants_fungi/ch04s04.html The walls of tracheid are moderately thick and show different types of cell wall thickening due to the deposition of secondary wall substances such as lignin. The bordered pits in tracheary elements show three main types of arrangement Scalariform pitting: Where elongated pits are arranged parallel to one another to form a ladder-like pattern, this pattern is called scalariform pitting. Opposite pitting: Circular or oval bordered pits arranged in horizontal pairs characterize opposite pitting. If such pits are crowded, their borders assume rectangular outlines in face view. Alternate pitting: When the pits are arranged in diagonal rows, this arrangement is called as alternate pitting, and their crowding results in borders that are Institute of Life Long Learning, University of Delhi 31 Plant Tissues polygonal (angular and with more than four sides) in outline and is most commonly found in dicots. Figure: Tracheids showing A) scalariform B) opposite and B) alternate pitting. Source: A) http://delta-intkey.com/wood/en/www/aquil-ho.htm B) http://blackwalnut.npust.edu.tw/archives/wood/133477; C) http://blackwalnut.npust.edu.tw/archives/wood/133478 Vessels Vessels or trachea are elongated tube like structure arranged in longitudinal series. The vessels are variable in length and its lengths are positively correlated with its diameters. Wide vessels are longer and narrow vessels are shorter. The longest vessels are found in the early wood of ring porous species of dicots in which the vessels (pores) of the firstformed wood (early wood) are especially wider. The large-diameter vessels have been found to extend throughout the entire length of tree’s stem but most of them were much shorter. The largest reported vessel element are approximately 3 meter in ash plant (Fraxinus excelsior; family Oleaceae), 18 meters in Fraxinus americana , 10.5 to 11.0 meters in Quercus rubra and smallest one is 60 cm found in Acer. Unlike tracheids, vessels are perforated at end walls except at the terminal end of vertical axis. The lumen of vessel is wider than that of tracheids. Vessel elements have perforations at both end through which vessel elements are interconnected. The imperforations are areas lacking both primary and secondary walls. The part of vessel Institute of Life Long Learning, University of Delhi 32 Plant Tissues element wall bearing the perforation or perforations is called perforation plate. The perforations also occur on the lateral wall. The vessel elements occur parallel to the long axis of plant body. The perforation plate may be simple or multiple one type. When due to dissolution of entire end wall, a single pore is formed at the perforation plate known as simple perforation plate (Mangifera). On the other hand, if the perforation plate has many pores, then it is called multiple perforation plate (Liriodendron). The presence of simple type plate is characteristic of advanced type of plants where dissolution of end wall is more or less complete. Figure: A) L.S. of vascular bundles in spurge (Euphorbia) showing simple perforation; B) Compound perforation plate Source: A) http://www.sbs.utexas.edu/mauseth/weblab/webchap7xylem/7.3-5a.htm B) http://www.sbs.utexas.edu/mauseth/weblab/webchap7xylem/7.3-7.htm The secondary wall thickenings of vessels elements are seen as annular, spiral, scalariform, reticulate or pitted as found in tracheids. Scalariform perforation plates are formed when the perforation is elongated and are arranged in parallel series. When it is formed in the form of network it is called as reticulate but when it is in circular form called as foraminate perforation. Institute of Life Long Learning, University of Delhi 33 Plant Tissues Figure: (A) Acer saccharum wood, T. S. showing prominent reticulate pitting, vessels, and multiseriate rays. (B) L. S. of corn stem (Zea mays) through a vascular bundle, with phloem on the right, xylem on the left where xylem vessel elements with annular secondary walls Source:(A)http://images.botany.org/set-17/17-117h.jpg (B)http://www.sbs.utexas.edu/mauseth/weblab/webchap7xylem/7.2-1.htm Vessels are principal water conducting elements in angiosperms. Vessels are completely absent in pteridophytes and gymnosperms (except Gnetum). The main function of vessel is conduction of water and minerals along with mechanical support to the plant. Xylem fibres The fibres associated with xylem are known as xylem fibres. Xylem fibres are very much elongated with tapering ends. The fibres are dead cells having lignified walls with narrow lumen. They are present both in primary and secondary xylem. Xylem fibres or wood fibre are two types a) Fibres tracheids b) Libriform fibres Institute of Life Long Learning, University of Delhi 34 Plant Tissues Fibre tracheids are intermediate forms between fibre and tracheids, possessing border pits whose borders are not fully developed. Libriform fibres are narrow, having obliterated lumen. They contain thick secondary wall with simple pits. They provide additional mechanical support to the plant body. Modification of fibre-tracheids and libriform fibres are seen as new form called as gelatinous fibres which are the common components of reaction wood in dicots. Xylem parenchyma Living parenchyma cells associated with the xylem are known as xylem parenchyma. It is the only living component found in xylem tissue. Xylem parenchyma cell wall is thin and made up of cellulose. They act as storage house of starch and fat with assisting in conduction of water. Two forms the xylem parenchyma cells are present in the secondary xylem: a) axial parenchyma b) ray parenchyma The axial parenchyma is the cells which are derived from the elongated fusiform initials of vascular cambium, and their long axes are oriented vertically in root or stem. The ray parenchyma cells are derived from the short ray initials of the vascular cambium. They may have their long axes oriented either vertically or horizontally with regard to the axis of stem or root. Figure: Showing axial and ray parenchyma in secondary xylem. Source:http://www.cas.miamioh.edu/~meicenrd/ANATOMY/Ch5_CellTypes/parenchyma. html Institute of Life Long Learning, University of Delhi 35 Plant Tissues When these two type of cells present in the secondary xylem there secondary walls are well lignified. The pit-pairs are present between parenchyma cells. They may be bordered, half-bordered, or simple but they are always simple. When some parenchyma cells deposit thick secondary walls they are known as sclerotic cells, or sclereids. The xylem parenchyma cells can store food reserves in the form of starch and fat. These content shows rhythmic fluctuation that help in adaptation. In many deciduous trees of temperate region, starch accumulates in late summer or early autumn but declines during dormancy period. It is because low winter temperature favors conversion of starch to sucrose which further acts as protecting agent during full dormancy against frost injury and starch is again resynthesized at the end of dormancy. Apart from fat and starch, tannins and crystals are also common inclusions. These storage products help in identification of woods. The ray parenchyma cells in herbaceous plants and young twigs of woody plants shows presence of chloroplasts. The axial and the ray parenchyma cells are located adjacent to the vessels in the secondary xylem. These parenchyma cells develop protrusions or balloon like structure that enters through the pit cavities and into vessels lumen known as tyloses (singular: tylose). The parenchyma cells that give rise to tyloses are called as contact cells. Contact cell wall are characterized by less cellulose content but rich in pectin and deposited by the protoplast after completion of secondary wall formation. This layer is called the protective layer which is deposited on surfaces of the contact cell wall but is thickest on the side of the cell bordering the vessel, especially at the pit membrane. In addition to secondary xylem, tyloses also occur in primary xylem. The formation of tyloses, results in ceased activity or functions of vessels. Institute of Life Long Learning, University of Delhi 36 Plant Tissues Figure: Showing tyloses in xylem vessels Source:http://wwwplb.ucdavis.edu/labs/rost/virtual%20grape%20dreamweaver/Shoots %20Secondary%203.html Phloem Phloem is also a complex tissue. It is the principal food-conducting tissue of vascular plants and it also transports diverse range of substances such as sugars, amino acids, micronutrients, lipids (primarily in the form of free fatty acids), hormones, floral stimulus (florigen), proteins and RNAs. Phloem plays an important role in inter-organ communication and coordination of growth processes within the plant. Long-distance signaling in plants occurs mainly through the phloem. Apart from food transport, a large amount of water is also transported through them which serve as the principal source of water for fruits, young leaves, and storage organs such as tubers. Phloem is spatially associated with the xylem. Phloem may be classified as primary or secondary on the basis of its time of appearance in relation to the development of the plant or organ as a whole. The primary phloem is derived from the procambium of apical meristem and initiated in embryo or young seedling is constantly added to during the development of the primary plant body and completes its differentiation when the primary plant body is fully formed. The secondary phloem originates from vascular cambium and reflects the organization of this meristem in its possession of axial and radial systems. The phloem rays are continuous through the cambium with those of xylem, providing a pathway for radial transport of substances between the two vascular tissues. The primary phloem elements that develop first from the procambium are smaller in size called the protophloem, whereas those develop later are larger in size called metaphloem. The protophloem is short lived. It is crushed by the developing metaphloem. Primary and secondary phloem tissues contain the same types of cells but the organization is different. The primary phloem is made up of one system without rays but secondary phloem is organized into the axial and the radial system with rays. Institute of Life Long Learning, University of Delhi 37 Plant Tissues Phloem is composed of four kinds of cells. a) Sieve elements b) Companion cells c) Phloem parenchyma d) Phloem fibres B A Figure: A) Showing components of phloem tissue B) Showing Sieve tube element, companion cell and sieve plate. Source:A)http://elte.prompt.hu/sites/default/files/tananyagok/plants_fungi/ch04s04.html B) http://preuniversity.grkraj.org/html/3_PLANT_ANATOMY.htm Apart from these four types of cells, fibres and sclereids are also common phloem components. Sometimes laticifers, resin ducts, and idioblasts may also be present in the phloem. Companion cells are present only in angiosperms and are absent in pteridophytes and gymnosperms. Phloem fibres are absent in primary phloem of most of angiosperms. But they are usually present in the secondary phloem. Sieve elements Sieve elements are main component of phloem because of the presence in their walls of areas (sieve areas) penetrated by pores. Sieve elements can be segregated on the basis of specialization into sieve tubes or sieve cells. Sieve tubes are more specialized but sieve Institute of Life Long Learning, University of Delhi 38 Plant Tissues cells are less specialized. The sieve tube forms a longitudinal series of sieve-tube elements similar to vessel elements. Contrary to the vessel elements, the distribution of sieve-tube elements is restricted only in angiosperms and use of term sieve cell is restricted to gymnosperms. The sieve elements of the seedless vascular plants, or vascular cryptogams, show much variation in structure and development and are referred as sieve element. Young sieve elements contain the entire cellular components characteristic of young plant cells. As they differentiate, the sieve elements undergo profound changes. Sieve tubes Sieve tubes are long tube like structure and are formed from a row of cells arranged in longitudinal series. The end wall of sieve tube cells shows sieve cell like perforation called as sieve plate which establishes the connectivity with the neighboring sieve cell. The protoplasts of sieve-tube elements contain P-protein (phloem protein or formerly called as slime). The unique carbohydrate known as callose is associated with conducting sievetube elements and it is deposited there in response to mechanical injury, some kind of stimulation. Callose normally accumulates at sieve plates and lateral areas and disappears sometime after the sieve element dies. Callose apparently plays a role in sieve-pore development. Figure: L.S. of stem showing components of phloem tissue Source:http://lurnq.com/lesson/Anatomy-of-Flowering-Plants-Part-I-Tissues/ Institute of Life Long Learning, University of Delhi 39 Plant Tissues The sieve-tube elements are associated with companion cells which is a specialized parenchyma cells. Thus it is called as sieve tube-companion cell complex or sieve element–companion cell complex. It is closely related to the sieve-tube elements both ontogenetically and functionally. Sieve cells The sieve cells are narrow elongated cells without conspicuous sieve areas. Sieve cells possess more sieve area. Sieve cells have sieve areas on their lateral walls only and are not arranged one above the other in linear rows. They are not associated with companion cells. They are more primitive than sieve tubes and distributed mainly in lower vascular plants and gymnosperms. Sieve cells have acted as precursor cell and give rise of sieve tubes. Companion cells These are specialized thin-walled, elongated parenchyma cells intimately associated with sieve-tube elements. Companion cells and sieve-tube elements are closely related ontogenetically and derived from the same mother cell. In contrast to sieve elements, the companion cell cytoplasm has a prominent nucleus and is connected to sieve tubes through pits found in lateral walls. The companion cells are present only in angiosperms and occur abundantly in monocots. Companion cells are absent in primitive dicots and in primary phloem. Gymnosperms and pteridophytes lack companion cell but have analogous cell known as albuminous cell. It is also known as Strasburger cell and named after Eduard Strasburger. The Strasburger cells show symplastic connections and have large median cavities containing numerous elements of smooth tubular endoplasmic reticulum with the sieve cells. In contrast to companion cells, the albuminous cells are smaller in size and have different origin. They assist the sieve tubes in the conduction of food materials. Phloem parenchyma The parenchyma cells associated with phloem are called phloem parenchyma. These are living cells with their cell wall rich in cellulose and primary pit fields. They are concerned with storage of starch, fats, resins and tannins. The parenchyma cell of primary phloem is elongated and is present with sieve elements along long axis. The parenchyma cell in secondary phloem are two types, phloem parenchyma and ray cells. Former type is Institute of Life Long Learning, University of Delhi 40 Plant Tissues associated with vertical plane and later one is associated with horizontal plane. Usually parenchyma cells are absent in monocots. Phloem fibres The fibres of sclerenchyma associated with phloem are called phloem fibres or bast fibres. They are narrow, vertically elongated cells with thick walls and a narrow lumen (the cell cavity). Among the four kinds of phloem elements, phloem fibres are the only dead tissue. Phloem fibres are rare in pteridophytes and some spermophytes but occur both in primary and secondary phloem. They functions as strengthening and supporting cells. Summary Tissues are made up of multiple cells having common origin and together perform a similar or a set of similar functions. These tissues perform various functions necessary to maintain biological life. Meristematic tissues have similar cells that have retained their capacity to divide and form new cells. These new cells lose the ability to form permanent tissue. Some plant tissues are made up of only one type of cells called as “simple” tissues whereas others are composed of different cell types called as “complex” tissue. Parenchyma is a versatile ground tissue that constitutes the “filler” tissue in soft parts of the plants. Parenchyma cells usually have thin primary walls. Parenchyma tissue is involved in almost all the physiological function of the plant. They are involved in the storage of food, provide turgidity to the softer parts and cortex of aquatic plant have parenchyma cells with large air spaces and cavities. Collenchyma is another group of simple tissue consisting of living cells found in plants. These are elongated, soft pliable cells with non-lignified thickened primary walls. Due to thick wall nature, it has specialized to function as supporting tissue. Cell wall of collenchyma contain in addition to cellulose, large amounts of pectin and hemicelluloses and without lignin deposition. On the basis of thickening on tangential wall, three types of collenchyma cells are found such as plate or lamellar, angular and lacunar/lacunate collenchyma. Sclerenchyma is a specialized tissue consisting of a group of cells in which secondary walls are lignified. On the basis of length, sclerenchyma cells have been kept in two Institute of Life Long Learning, University of Delhi 41 Plant Tissues groups; fibres and sclereids. Usually the fibre cells are long while sclereids are short cells. Angiosperms contain fibres in stem as outermost part of primary phloem as anastomosing strands or tangential plates. On the basis of position of fibre in plant body, they have been kept in two groups, xylary or intraxylary fibre and extraxylary fibres. Fibres present in xylem are called xylary. Extarxylary fibres are located outside the xylem and these are of three types; phloem fibre, pericyclic or perivascular fibre and cortical fibre. Sclereids are type of sclerenchyma cells. They are short cells having thick and strongly lignified secondary walls with many simple pits. The secondary walls are multilayered. On the basis of size and shape of sclereids, they have been classified into macrosclereids, brachysclereids, osteosclereids, astrosclereids and trichosclereids. A tissue that consists of several kinds of cells but all of them function together is called complex tissue. Xylem and phloem are example of complex tissue. They are involved in the water-conduction, transport of solutes and food material, are known as vascular tissue. Two type of xylem tissue have been demarcated in plants - primary xylem and secondary xylem. Xylem tissue is consists of four kinds of cells namely tracheids, vessels or tracheae, xylem fibres and xylem parenchyma. Phloem is also a complex tissue and second important tissue present in vascular tissue. Phloem is composed of four kinds of cells, sieve elements, companion cells, phloem parenchyma and phloem fibres. The phloem is the principal food-conducting tissue of vascular plants. Glossary Angular collenchyma: It is a form of collenchymas in which the primary wall thickening is most prominent in the angles where several cells are associated. Astrosclereid: It is a type of sclereid in which shows branching or ramification. Brachysclereid: It is also known as stone cell. It is short, roughly isodiametric sclereid and resembling a parenchyma cell in shape. Collenchyma: It is supporting tissue composed of more or less elongated living cells with unevenly thickened, nonlignified primary walls. Collenchyma is prominant in regions of primary growth in stems and leaves. Institute of Life Long Learning, University of Delhi 42 Plant Tissues Companion cell: A specialized parenchyma cell associated with a sieve-tube element in angiosperm phloem and arising from the same mother cell as the sieve-tube element. Lacunar collenchyma: A type of collenchymas tissue characterized by intercellular spaces and cell wall thickenings facing the intercellular spaces. Lamella: A thin plate or layer. Lamellar: A type of collenchymas tissue in which cell wall thickenings are deposited mainly on tangential walls. Macrosclereid: It is a type of sclereids which is elongated with unevenly distributed secondary wall thickening and commonly found in seed epidermis of Fabaceae. Metaphloem: It is a part of the primary phloem that differentiates after the protophloem and before the secondary phloem. Metaxylem: It is a part of the primary xylem that differentiates after the protoxylem and before the secondary xylem. Osteosclereid: It is a type of sclereids which is Bone-shaped and having a columnar middle part with enlargements at both ends. Parenchyma cell: Generally it is not a specialized cell having a nucleate protoplast and associated with one or more of the various physiological and biochemical activities in plants. Parenchyma cells are diverse in size, form, and wall structure. Phloem parenchyma: A type of parenchyma cells located in the phloem are referred as axial parenchyma in secondary phloem. Phloem: It is main food-conducting tissue of the vascular plant and contains sieve elements phloem parenchyma cells, fibres and sclereids. P-protein: It is a proteinaceous material found in Phloem especially sieve tube elements. Institute of Life Long Learning, University of Delhi 43 Plant Tissues Primary phloem: When Procambium differentiates it give rise of phloem tissue during primary growth and differentiation of a vascular plant. It is then commonly divided into the earlier protophloem and the later metaphloem. Protoxylem: It is the First-formed elements of the xylem. Ray parenchyma: It is a type of parenchyma cells found in secondary vascular tissues. Sclereid: It is a sclerenchyma cell, short and thick lignified secondary walls with many pits. Sclerenchyma: It is a type of tissue composed of sclerenchyma cells that includes fibres, fibresclereids, and sclereids. Sieve cell: It is a type of sieve element that has relatively undifferentiated sieve areas without sieve plates. Generally sieve cells are found in phloem of gymnosperms. Sieve element: These are cell in the phloem tissue concerned with mainly longitudinal conduction of food materials. Sieve plate: It is a part of the wall of a sieve-tube element bearing one (simple sieve plate) or more (compound sieve plate) highly differentiated sieve areas. Sieve tube: A series of sieve-tube elements arranged end to end and interconnected through sieve plates. Sieve-tube element: It is one of the series of cellular components of a sieve tube. It shows a more or less pronounced differentiation between sieve plates (wide pores) and lateral sieve areas (narrow pores). Strasburger cell: Few ray and axial parenchyma cells spatially are functionally associated with the sieve cells resembling the companion cells of angiosperms. These do not have common origin from the same precursory cells as seen in the sieve cells and companion cells. They are also known as albuminous cells and are mainly associated with gymnosperm phloem. Institute of Life Long Learning, University of Delhi 44 Plant Tissues Tissue: It is a group of cells organized into a structural and functional unit. Tissue system: Structurally and functionally organized tissue or tissues in a plant or plant organ into a unit is called as tissue system. In plants, commonly three tissue systems are recognized, dermal, vascular and fundamental (ground tissue system). Tracheary element: water conducting cell, tracheid or vessel element found in vascular system of plants. Tracheid: A tracheary element of the xylem. It has no perforations as found in vessel element and may have any kind of secondary wall thickening. Tracheids occur in primary and in secondary xylem. Trichosclereid: It is type of branched sclereid with hair-like branches extending into intercellular spaces. Tylose (pl. tyloses): It is an outgrowth from found in xylem where a parenchyma cell (axial or one in a ray) through a pit cavity into a tracheary cell and partially or completely blocking the lumen of tracheids. Vascular tissue: It refers to either or both vascular tissues, xylem and phloem. Vessel: It is a tube-like series of vessel elements and the common walls of have perforations. Vessel element: It is one of the cellular components of a vessel. Xylem elements: It is a cell composing the xylem tissue. Xylem fibre: A fibre of the xylem tissue. Xylem: It is a principal water-conducting tissue in vascular plants characterized by the presence of tracheary elements. Xylem ray: Part of a vascular ray that is located in the secondary xylem. Institute of Life Long Learning, University of Delhi 45 Plant Tissues Exercises Q.1. Distinguish between following pairs: a) Paranchyma and Collenchyma tissue b) Parenchyma and fibre cells c) Xylem and phloem tissue d) Companion and albuminous cell e) Macrosclereids and osteosclereids f) Bast fibre and surface fibre g) Brachysclereids and Trichosclereids h) Angular and lacunar collenchyma i) Cortical and Phloem fibre j) Intraxylary and extraxylary fibre Q. 2. Write short note on following topics: a) Trachieary elements b) Sieve tube c) Trichosclerids d) Strasburger cell e) Sclereids f) Structural fibre g) Sieve Cells h) Phloem fibre Q.3. Describe the general properties of parenchyma cells. Q.4. What is sclereids and write its significance. Q.5. Describe the type of tissue found in in angiosperms. Q.6. Explain the collenchyma and write its importance in plant system. Q.7. List the types of parenchyma tissue with examples. Q.8. Discuss the types of fibre with suitable example. Institute of Life Long Learning, University of Delhi 46 Plant Tissues Q.9. Discuss the anatomical difference between tracheid and vessel of vascular plants. Q.10. What is xylem parenchyma and write its function. Q.11. Define collenchymas and describe the types of collenchyma with suitable examples. Q.12. what is vascular tissue. Elaborate their components with their function. Multiple Choice Questions Q.1: Collenchymatous hypodermis is characteristic feature of (a) Dicot stems (b) Grass Stem (c) Monocot stem (d) Plant roots Correct Answer: (a) Dicot stem Feedback for answer: Collenchyma is present in the hypodermal region of dicot stem providing mechanical support. Resource/Hint/feedback for the wrong answer Collenchyma is absent from roots and monocot plants Q.2: Who gave the term tissue (a) Hooke (b) Charles Darwin (c) Nageli (d) N. Grew Institute of Life Long Learning, University of Delhi 47 Plant Tissues Correct Answer: (d) Grew Feedback for answer: N. Grew coined the term tissue in 1682. Resource/Hint/feedback for the wrong answer Charles Darwin is known for many contributions in biology such as theory of evolution, natural selection and origin of species. Nageli coined the term meristem in 1858. Hooke discovered the cell in cork slice via rudimentary microscope. Q.3: A group of isodiametric cells with prominent intercellular space is called as (a) Collenchyma (b) Paranchyma (c) Chlorenchyma (d) sclerenchyma Correct Answer: (b) Paranchyma Feedback for answer: Paranchyma cells are basic type of cells are usually isodiamentric having 14 facets with intercellular space. Resource/Hint/feedback for the wrong answer Chlorenchyma cells are modified parenchyma cells which have developed chlorophyll and found in photosynthetic mesophyll cell. Sclerenchyma cells are dead cells with thick lignifications meant for mechanical function in plants. Collenchyma cells are living cells with high deposition of cellulose and pectin at the corner of cells. Institute of Life Long Learning, University of Delhi 48 Plant Tissues Q.4: A simple mechanical tissue rich in lignin is (a) parenchyma (b) sclerenchyma (c) collenchyma (d) chlorenchyma Correct Answer: (b) sclerenchyma Feedback for answer: Sclerenchyma cells are dead cells with thick lignifications meant for mechanical function in plants. Resource/Hint/feedback for the wrong answer Collenchyma is living cells with high deposition of cellulose and pectin at the corner of cells. Parenchyma cells are isodiametric cells with prominent intercellular space. Chlorenchyma cells are modified parenchyma cells which have developed chlorophyll and found in photosynthetic mesophyll cell. Q.5: Vascular bundles in a dicot stem are (a) open, collateral and exarch (b) closed, collateral and endarch (c) closed, collateral and exarch (d) open, collateral and endarch Correct Answer: (d) open, collateral and endarch Institute of Life Long Learning, University of Delhi 49 Plant Tissues Feedback for answer: Dicot stem shows the presence of conjoint, collateral and open vascular bundles with an endarch xylem. Resource/Hint/feedback for the wrong answer (a) Dicot stem vascular bundles are not open, collateral and exarch type (b) Dicot stem vascular bundles are not closed, collateral and endarch (c) Dicot stem vascular bundles are not closed, collateral and exarch Q.6: Sieve tubes are suited for translocation of food because they possess (a) bordered pits (b) no ends walls (c) broader lumen and perforated cross walls (d) no protoplasm. Correct Answer: (c) Broader lumen and perforated cross walls. Feedback for answer: Sieve tubes are suited for translocation of food because they possess broader lumen and perforated cross walls. Resource/Hint/feedback for the wrong answer Border pits are absent They posses end walls known as sieve plates They have protoplasm that lacks nuclei at maturity. Q.7: Angular collenchyma occurs in (a) Cucurbita Institute of Life Long Learning, University of Delhi 50 Plant Tissues (b) Helianthus (c) Ipomoea (d) Sombucus Correct Answer: a) Cucurbita Feedback for answer: Angular collenchyma occurs in Cucurbita. It has thickening at the angles and there are no intercellular spaces. It is generally found in leaf petioles. Resource/Hint/feedback for the wrong answer Helianthus shows lamellar collenchymas in their stem Sambucus stem shows lamellar collenchymas in their stem Ipomoea stem shows lacunar collenchyma in their stem Q.8: Bordered pits are found in (a) Sieve cells (b) Vessel wall (c) Companion cells (d) Sieve tube wall. Correct Answer: (b) Vessel wall Feedback for answer: Bordered pits are cavities in the lignified cell walls of xylem conduits (vessels and tracheids) that are essential components in the water-transport system of higher plants. Institute of Life Long Learning, University of Delhi 51 Plant Tissues Resource/Hint/feedback for the wrong answer Sieve cells are long, slender, conducting cells of the phloem don’t have bordered pits. Companion cell are living cell associated with sieve tube cells in phloem in angiosperms and don’t possess bordered pits. Sieve tube wall lack bordered pits. Q.9: Which of the following statement is not true about ‘sclereids’?? (a) These are groups of living cells. (b) These are found in nut shells, guava pulp, pear. (c) These are also known as stone cells. (d) These are form of sclerenchyma with fibres. Correct Answer: (a) Sclereids are a type of sclerenchyma cells. Feedback for answer: Sclereids are a reduced form of sclerenchyma cells with highly thickened, lignified cellular walls Resource/Hint/feedback for the wrong answer Sclerieds are found in nut shells, guava pulp, pear. These are alternatively known as stone cells Sclerieds are form of sclerenchyma with fibres. Institute of Life Long Learning, University of Delhi 52 Plant Tissues Q.10: Vessels are reported in (a) All angiosperms and few gymnosperms (b) Most of angiosperms and few gymnosperms (c) All angiosperms, all gymnosperms and few pteridophyta (d) all pteridophyta and bryophyta Correct Answer: (b) Most of angiosperms and few gymnosperms Feedback for answer: Vessels are long tubelike nonliving lignified structures meant for the conduction of water and solutes. Resource/Hint/feedback for the wrong answer Vessels are found in the wood of almost all the angiosperms except certain primitive members of the order ranales (vesseless dicots), e.g., Trochodendron, Tetracentron, Drimys, Pseudowintera, etc. Vessels also occur in the members of order Gnetales of gymnosperms (e.g., Genetum, Ephedra and Welwitschia) and in some pteridophytes. References 1. Esau, K. 1965. Plant Anatomy. Jhon Wiley and Sons Inc., New York. 2. Ray F. Evert. 2006. Esau's Plant Anatomy: Meristems, Cells, and Tissues of the Plant Body: Their Structure, Function, and Development, Third Edition. Jhon Wiley and Sons Inc., New York. Institute of Life Long Learning, University of Delhi 53 Plant Tissues 3. Fahn, A. 1990. Plant Anatomy. Pergamon Press, Oxford. 4. William C. Dickison. 2000. Integrative Plant Anatomy, Academic Press. 5. V. Singh, P.C. Pande, D.K. Jain. 2005. Anatomy of Seed Plants, Rastogi Publications, Meerut. 6. 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