The cell wall provides structure and protection for plant cells. It has three main layers - the middle lamella, primary wall, and secondary wall. The middle lamella binds adjacent cells, the primary wall is the first layer deposited and allows cell growth, and the secondary wall provides support and is thicker with lignin. The cell wall is composed of cellulose microfibrils in a matrix of pectin and hemicellulose. It also contains structural components like lignin, cutin and suberin. Pit pairs and plasmodesmata allow communication between cells and transport through the cell wall. Bordered and simple pits are thin areas in the secondary wall that facilitate water and nutrient transport.
5. Primary wall First wall the develops on new cell Cellulose, pecticcpds., non-cellulosic polysaccharides and hemicellulose May be lignified Assoc. with living protoplasts --eg. meristematic cells, parenchyma, collenchyma
6. Secondary wall Formed in the inner surface of P-wall Same content as Pwall ( > cellulose) + lignin In cells that ceased to grow; devoid of protoplast at maturity * xylem ray, xylem parenchyma – still living Mechanical support Compound middle lamella * = 3-layered or 5-layered = middle lamella + 2 P-walls (+ 2 S-walls) *if middle lamella is obscured
8. Cell plate- precursor of cell wall; rich in pectins Phragmoplast- a complex of microtubules and ER that forms during late anaphase or early telophase from dissociated spindle subunits.
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10. Fine structure of the wall Cellulose fibrils Matrix (non-cellulosic): - with lignin, cutin, suberin, hemicelluloses etc.
14. Levels of organization of cell wall Long chains of linked glucose residues Micellae – bundles of cellulose molecules or ELEMENTARY FIBRIL = ~40 cellulose molecules Microfibril Bundles of microfibril
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16. CHEMISTRY OF WALLS Cellulose Pectic substances Gums and mucilages Lignin Fatty substances
19. Gums and mucilages Appear as a result of physiological or pathological disturbances that induce breakdown of walls and cell contents
20. Lignin Phenolic compounds May be found in middle lamella, primary wall, and secondary wall hydrophobic filler that replaces the wall’s water compressive strength and bending stiffness Microbial attack resistance
21. Fatty substances Cutin, suberin, waxes Waxes- glaucous condition; assoc. with cutin and suberin Suberin- cork cells of periderm; endodermis and exodermis; prevents apoplastic transport Cutin- cuticle layer; epidermis of aerial parts Cutinization, suberization- impregnation in cell wall Cuticularization- formation of layer
22. Cellulose Tensile strength (bend under compressive stress) Incrustation– eg. Lignification Cell wall growth A. intussusception B. apposition C. mosaic growth D. multinet growth
23. Intussusception Material of new wall is laid down bet. Particles of the existing substance of the expanding wall
24. Apposition Growth is due to the centripetal addition of new layers one upon the other
25. Mosaic growth Fibrillar texture in certain wall areas become loosened as a result of turgor pressure and afterwards mended by deposition of new microfibrils in the gaps caused by the strain
26. Multinet growth separation of crossed microfibrils and alteration in their orientation transverse longitudinal
27. Special structures of the cell wall Primary pit fields Pits Crassulae Trabeculae Wart structures Cystoliths
28. Primary pit fields Primordial pits/ primary pit fields Certain areas of primary wall of young cells remain thin May appear beaded in xs
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31. Plasmodesmata Plasmodesmata- connnect protoplasts of neighboring cells - transport; relay of stimuli * symplast- 2 or more interconnected protopolast * apoplast – cell walls, intercellular spaces and lumen desmotubule
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35. Pits Portions of the cell wall that remained thin even as secondary wall is formed Primary wall only Can develop over primary pit fields Function?
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37. Pits TYPES: a. simple pit b. bordered pit—S-wall develops over the pit cavity to form an overarching roof
38. Simple pits Branched simple pits (ramiform) Found in parenchyma cells with thickened walls, libriform fibers, sclereids, phloem fibers
47. Torus pit membrane thickening; disc shaped flexible; can go median or lateral Aspirate condition (lateral)– latewood and all heartwood Coniferales, Gnetales
48. Margo – porous pit membrane around the torus --conifer tracheids -- occurs through matrix dissolution
49. Shape of pit aperture Round, elliptic, linear In thick cell walls: *inner aperture becomes long and narrow *outer aperture remains circular round * pit canal is funnel-shaped *fiber-tracheid feature
52. crassulae Linear or crescent-shaped thickenings of the primary wall and middle lamella gymnosperms
53. trabeculae Rod shape thickenings of the wall which traverse the cell lumen radially
54. References Fahn, A. 1990. Plant Anatomy, 4th ed.. Pergamon Press Esau, K. 1958. Plant Anatomy. John Wiley and Sons, Inc. Evert, R. 2006. Esau’s Plant Anatomy. John Wiley and Sons, Inc.
Notas do Editor
(A) In the first step, Golgi vesicles, some of which are interconnected via fusion tubes, aggregate in the spindle midzone area. This structure is called the fusion tube network (FTN). The transition from the first to the second stage of cell plate formation can be inhibited by caffeine. (B) Formation of a tubulo-vesicular network (TVN). The contents of the vesicles, mainly pectins, represent the precursors from which the new middle lamella is assembled outside the cell. In the next stage, vesicle fusion increases, forming a tubulo-vesicular network (TVN), and the membranes become coated with either clathrin or other proteins. (C) In the third stage, the central region of the growing cell plate forms a tubular network (TN), with vesicle fusion occurring at the growing edges where the remaining microtubules are located. (D) In the final stage, the cell plate contacts and adheres to the plasma membrane of the parent cell. At the same time the tubular network expands to form a fenestrated sheet. (E) At the end of mitosis, the phragmoplast disappears, the cell enters interphase, and microtubules reappear in the cytosol near the plasma membrane, where they play a role in the deposition of cellulose microfibrils during cell wall growth (see Chapter 15). (F) Electron micrograph of a cell plate forming in a root tip of a beet, (Beta vulgaris) (10,000×) MT, microtubule; VE, vesicles; N, nucleus; NE, nuclear envelope; P, cell plate. (A–E from Staehelin and Hepler 1996; F from B. Gunning and M. Steer, Plant Cell Biology: Structure and Function, Jones and Bartlett, 1996.)Stages of cell plate development. A, the fusion of Golgi-derived secretory vesicles (sv) at the equatorial zone, among phragmoplast microtubules (mt) and a cytoplasmic fuzzy matrix (fm). B, fused Golgi-derived vesicles give rise to tubulo-vesicular network covered by a “fuzzy coat.” C, a tubular network (TN) forms as the lumen of the tubulovesicularnetwork (TVN) becomes filled with cell wall polysaccharides, especially callose. Fuzzy matrix surrounding the network and microtubules disappears, further distinguishing this stage from the tubulo-vesicular network. D,the tubule areas expand, forming an almost continuous sheet. Numerous finger-like projections extend from the margins of the cell plate and fuse with the plasma membrane (pm) of the parent cell wall (pcw) at the site previously occupied by the preprophase band. E, maturation of the cell plate into a new cell wall. (After Samuels et al., 1995.Reproduced from The Journal of Cell Biology 1995, vol. 130, 1345–1357, by copyright permission of the RockefellerUniversity Press.)
Primary plasmodesmata form during cytokinesis when Golgi-derived vesicles containing cell wall precursors fuse to form the cell plate (the future middle lamella). Rather than forming a continuous uninterrupted sheet, the newly deposited cell plate is penetrated by numerous pores (Figure 1.27A), where remnants of the spindle apparatus, consisting of ER and microtubules, disrupt vesicle fusion.Further deposition of wall polymers increases the thickness of the two primary cell walls on either side of the middle lamella, generating linear membrane-lined channels (Figure 1.27B). Development of primary plasmodesmata thus provides direct continuity and communication betweencells that are clonally related (i.e., derived from the samemother cell)