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1/13/2010




                                                Gross Structure
                                                Detailed Structure
                                                Chemistry
                                                Features




                                                Cement; amorphous subs.
                                                Bet. P‐walls of neighboring cells
                                                Pectic substances (Ca, Mg pectate)




First wall the develops on new cell             Formed in the inner surface of P‐wall
                                                Same content as Pwall ( > cellulose) + lignin
Cellulose, pectic cpds., non‐cellulosic         In cells that ceased to grow; devoid of protoplast at 
                                                maturity
polysaccharides and hemicellulose               * xylem ray, xylem parenchyma – still living
May be lignified
      b l    f d                                Mechanical support
                                                M h i l 
Assoc. with living protoplasts
‐‐eg. meristematic cells, parenchyma,      Compound middle lamella *
                                           = 3‐layered or 5‐layered
collenchyma                                = middle lamella + 2 P‐walls (+ 2 S‐walls)


                                           *if middle lamella is obscured




                                                                                                                1
1/13/2010




                                                       Preprophase band
                                                         a cortical belt of microtubules and actin filaments,
                                                       that predicts the plane of the future cell plate

                                                       Phragmosome
                                                       Ph
                                                         a layer of cytoplasm which spreads across the future 
                                                       division
                                                          contains microtubules and actin filaments




                                                       (1) the arrival of Golgi‐derivedvesicles in the 
                                                       division plane; fusion tube starts to form




                                                                Start of accumulation of cell wall materials in the lumen‐‐callose




(2) the formation of fusion tubes that grow out of    (3) transformation of the tubulo‐vesicular network 
  the vesicles and fuse with others tubulo‐             into a tubular network
  vesicular network (interwoven)




                                                                                                                                            2
1/13/2010




(4) Formation of fenestrated plate‐like structure             (5)the formation of numerous finger‐like projections 
  the dense membrane coat and the associated                    at the margins of the cell plate that fuse with the 
  phragmoplast microtubules are  disassembled;                  plasma membrane of the mother cell wall




     Abundant accummulation of cell wall building materials




(6) maturation of the cell plate into a new cell wall.


                                                                  Cell plate‐ precursor of cell wall; rich in 
                                                                  pectins

                                                                  Phragmoplast‐ a complex of microtubules 
                                                                   h         l           l   f         b l
                                                                  and ER that forms during late anaphase or 
                                                                  early telophase from dissociated spindle 
  Closing of fenestrae                                            subunits.
  Formation of plasmodesmata




                                                                  After completion of the cell plate, additional wall 
                                                                  material is deposited inc. in thickness
                                                                  New wall material deposition  mosaic fasion
                                                                  Matrix materials 
                                                                  Matri  materials      Delivered by Golgi vesicles
                                                                  Cellulose microfibrils Cellulose synthase complex
                                                                  Cellulose synthase
                                                                    appear as rosettes
                                                                    exude the microfibrils on the outer surface of 
                                                                  the membrane.




                                                                                                                             3
1/13/2010




                                                                                A. Growth in thickness
                                                                                  1. Apposition
                                                                                  2. Intussusception
                                                                                Lignification
                                                                                      f            Intussusception

                                                                                Cutinization       Intussusception



                                                                                B. Growth in surface (wall expansion)

 Rosettes are inserted in the plasma membrane and pushed forward by synthesis
 and crystallization of microfibrils




 Or extension
 Requires: wall stress relaxation (loosening of 
 wall structure) and turgor pressure
 Controlled by: a] amt. of turgor pressure
         ll d b           f
                 b] extensibility
*Extensibility  ability to expand permanently 
 when a force is applied to it (plastic)
                affected by hormones : auxin




   Marks cessation of growth (irreversible)                                       Cellulose fibrils
   during maturation                                                              Matrix (non‐cellulosic):
FACTORS: that contribute                                                          ‐ with lignin, cutin, suberin, hemicelluloses 
(1) a reduction in wall‐loosening processes, 
        d             ll l                                                        etc. 
(2) An increase in cross‐linking of cell wall 
   components,
 (3) a change in wall composition (more
rigid structure or less susceptible to wall 
   loosening)




                                                                                                                                          4
1/13/2010




                       Long chains of linked glucose residues
                       Micellae – bundles of cellulose molecules or 
                       ELEMENTARY FIBRIL =  ~40 cellulose 
                       molecules
                          l l
                       Microfibril
                       Bundles of microfibril
Ml‐ middle lamella
Pm‐ plasma membrance




                       Cellulose
                       Pectic substances
                       Gums and mucilages
                       Lignin
                       Fatty substances




                                                                              5
1/13/2010




  Hydrophilic crystalline compound                  Amorphous colloidal substances
  Repeating monomers of glucose                     Plastic and hydrophilic




  Appear  as a result of physiological or           Phenolic compounds
  pathological disturbances that induce             May be found in middle lamella, primary wall, 
  breakdown of walls and cell contents              and secondary wall
                                                    hydrophobic fi ller h
                                                    h d h b f ll that replaces the wall’s 
                                                                               l    h     ll
                                                    water
                                                    compressive
                                                    strength and bending stiffness
                                                    Microbial attack resistance




  Cutin, suberin, waxes                             Tensile strength  (bend under compressive 
  Waxes‐ glaucous condition; assoc. with cutin      stress)
  and suberin                                       Incrustation– eg. Lignification
  Suberin‐ cork cells of periderm; endodermis 
  and exodermis; prevents apoplastic transport
  Cutin‐ cuticle layer; epidermis of aerial parts   Cell wall growth
                                                    A. intussusception
Cutinization, suberization‐ impregnation in cell    B. apposition
 wall                                               C. mosaic growth
Cuticularization‐ formation of layer                D. multinet growth




                                                                                                        6
1/13/2010




  Material of new wall is laid down bet.            Growth is due to the centripetal addition of 
  Particles of the existing substance of the        new layers one upon the other
  expanding wall




  Fibrillar texture in certain wall areas become    separation of crossed microfibrils and 
  loosened as a result of turgor pressure and       alteration in their orientation
  afterwards mended by deposition of new            transverse  longitudinal
  microfibrils in the gaps caused by the strain
         fb l      h              db h




Primary pit fields                                  Primordial pits/ primary pit fields
Pits                                                Certain areas of primary wall of young cells 
Crassulae                                           remain thin
Trabeculae
   b l                                              May appear beaded in xs
                                                                b d d
Wart structures
Cystoliths




                                                                                                           7
1/13/2010




Plasmodesmata‐ connnect protoplasts of 
neighboring cells
‐ transport; relay of stimuli
* symplast‐ 2 or more interconnected 
        l                                d
protopolast
* apoplast – cell walls, intercellular spaces 
and lumen

desmotubule




                                                        8
1/13/2010




Portions of the cell wall that remained thin  
even as secondary wall is formed
Primary wall only
Can develop over primary pit fields
                                f

Function?




TYPES:                                                Branched simple pits (ramiform)
a. simple pit                                         Found in parenchyma cells with thickened 
b. bordered pit—S‐wall develops over the pit          walls, libriform fibers, sclereids, phloem fibers
cavity to form an overarching roof
          f              h       f




Structure:
 Pit cavity / pit chamber
 Pit aperture
 Pit border
 Pit canal, inner and outer aperture (very thick S‐
 wall)
water‐conducting and mechanical xylem cells 
(vessel elements, tracheids, etc.)




                                                                                                             9
1/13/2010




   Angiosperm                Gymnosperm




Pit cavity – break in S‐wall               Simple pit pair
Pit membrane/ closing membrane –primary    Bordered pit pair
wall + middle lamella                      Half bordered pit pair
Pit aperture                               Blind pit
                                            l d
                                           Unilateral compound pitting 




                                                                                10
1/13/2010




pit membrane thickening; disc shaped               – porous pit membrane around the torus
flexible; can go median or lateral                 ‐‐conifer tracheids
Aspirate condition  (lateral)– latewood and all    ‐‐ occurs through  matrix dissolution
heartwood
Coniferales, Gnetales




Round, elliptic, linear
In thick cell walls: 
*inner aperture becomes long and narrow
*outer aperture remains circular round
                             l       d
* pit canal is funnel‐shaped
*fiber‐tracheid feature




Scalariform                                        Linear or crescent‐shaped thickenings  of the 
Opposite                                           primary wall and middle lamella
Alternate                                          gymnosperms




                                                                                                      11
1/13/2010




Rod shape thickenings of the wall which      Fahn, A. 1990. Plant Anatomy, 4th ed.. 
traverse the cell lumen radially             Pergamon Press
                                             Esau, K. 1958. Plant Anatomy. John Wiley and 
                                             Sons, Inc.
                                             Evert, R. 2006. Esau’s Plant Anatomy. John 
                                             Wiley and Sons, Inc.




Protopectin, pectin, pectic acid             Related to pectic
Plastic, amorphous colloid, hydrophilic      Swelling property (hydrophilic)
Hydration in young walls




Impregnation starts in the intercellular    Cellulose molecules  micelles (crystal) 
lamella                                      micellar system (porous)

                                            Microfibril
                                                 f          consists of micellar system
                                                                      f
                                            And
                                            Microcapillaries liquids, lignin, waxes, cutin, 
                                             suberin, hemicellulose, pectic substances, 
                                             crystals, silica




                                                                                                 12
1/13/2010




Plasticity 
‐becoming permanently deformed when 
subjected to changes in shape or size
Elasticity
‐property of recovery of the original size and 
shape after deformation
Tensile strength
   ‐ relative to chemical composition and 
microscopic and submicroscopic structure




                                                        13

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Cell Plate Formation and Cell Wall Development

  • 1. 1/13/2010 Gross Structure Detailed Structure Chemistry Features Cement; amorphous subs. Bet. P‐walls of neighboring cells Pectic substances (Ca, Mg pectate) First wall the develops on new cell Formed in the inner surface of P‐wall Same content as Pwall ( > cellulose) + lignin Cellulose, pectic cpds., non‐cellulosic  In cells that ceased to grow; devoid of protoplast at  maturity polysaccharides and hemicellulose * xylem ray, xylem parenchyma – still living May be lignified b l f d Mechanical support M h i l  Assoc. with living protoplasts ‐‐eg. meristematic cells, parenchyma,  Compound middle lamella * = 3‐layered or 5‐layered collenchyma = middle lamella + 2 P‐walls (+ 2 S‐walls) *if middle lamella is obscured 1
  • 2. 1/13/2010 Preprophase band a cortical belt of microtubules and actin filaments, that predicts the plane of the future cell plate Phragmosome Ph a layer of cytoplasm which spreads across the future  division contains microtubules and actin filaments (1) the arrival of Golgi‐derivedvesicles in the  division plane; fusion tube starts to form Start of accumulation of cell wall materials in the lumen‐‐callose (2) the formation of fusion tubes that grow out of  (3) transformation of the tubulo‐vesicular network  the vesicles and fuse with others tubulo‐ into a tubular network vesicular network (interwoven) 2
  • 3. 1/13/2010 (4) Formation of fenestrated plate‐like structure (5)the formation of numerous finger‐like projections  the dense membrane coat and the associated  at the margins of the cell plate that fuse with the  phragmoplast microtubules are  disassembled; plasma membrane of the mother cell wall Abundant accummulation of cell wall building materials (6) maturation of the cell plate into a new cell wall. Cell plate‐ precursor of cell wall; rich in  pectins Phragmoplast‐ a complex of microtubules  h l l f b l and ER that forms during late anaphase or  early telophase from dissociated spindle  Closing of fenestrae subunits. Formation of plasmodesmata After completion of the cell plate, additional wall  material is deposited inc. in thickness New wall material deposition  mosaic fasion Matrix materials  Matri  materials  Delivered by Golgi vesicles Cellulose microfibrils Cellulose synthase complex Cellulose synthase appear as rosettes exude the microfibrils on the outer surface of  the membrane. 3
  • 4. 1/13/2010 A. Growth in thickness 1. Apposition 2. Intussusception Lignification f Intussusception Cutinization Intussusception B. Growth in surface (wall expansion) Rosettes are inserted in the plasma membrane and pushed forward by synthesis and crystallization of microfibrils Or extension Requires: wall stress relaxation (loosening of  wall structure) and turgor pressure Controlled by: a] amt. of turgor pressure ll d b f b] extensibility *Extensibility  ability to expand permanently  when a force is applied to it (plastic) affected by hormones : auxin Marks cessation of growth (irreversible)  Cellulose fibrils during maturation Matrix (non‐cellulosic): FACTORS: that contribute ‐ with lignin, cutin, suberin, hemicelluloses  (1) a reduction in wall‐loosening processes,  d ll l etc.  (2) An increase in cross‐linking of cell wall  components, (3) a change in wall composition (more rigid structure or less susceptible to wall  loosening) 4
  • 5. 1/13/2010 Long chains of linked glucose residues Micellae – bundles of cellulose molecules or  ELEMENTARY FIBRIL =  ~40 cellulose  molecules l l Microfibril Bundles of microfibril Ml‐ middle lamella Pm‐ plasma membrance Cellulose Pectic substances Gums and mucilages Lignin Fatty substances 5
  • 6. 1/13/2010 Hydrophilic crystalline compound Amorphous colloidal substances Repeating monomers of glucose Plastic and hydrophilic Appear  as a result of physiological or  Phenolic compounds pathological disturbances that induce  May be found in middle lamella, primary wall,  breakdown of walls and cell contents and secondary wall hydrophobic fi ller h h d h b f ll that replaces the wall’s  l h ll water compressive strength and bending stiffness Microbial attack resistance Cutin, suberin, waxes Tensile strength  (bend under compressive  Waxes‐ glaucous condition; assoc. with cutin stress) and suberin Incrustation– eg. Lignification Suberin‐ cork cells of periderm; endodermis  and exodermis; prevents apoplastic transport Cutin‐ cuticle layer; epidermis of aerial parts Cell wall growth A. intussusception Cutinization, suberization‐ impregnation in cell  B. apposition wall C. mosaic growth Cuticularization‐ formation of layer D. multinet growth 6
  • 7. 1/13/2010 Material of new wall is laid down bet.  Growth is due to the centripetal addition of  Particles of the existing substance of the  new layers one upon the other expanding wall Fibrillar texture in certain wall areas become  separation of crossed microfibrils and  loosened as a result of turgor pressure and  alteration in their orientation afterwards mended by deposition of new  transverse  longitudinal microfibrils in the gaps caused by the strain fb l h db h Primary pit fields Primordial pits/ primary pit fields Pits Certain areas of primary wall of young cells  Crassulae remain thin Trabeculae b l May appear beaded in xs b d d Wart structures Cystoliths 7
  • 8. 1/13/2010 Plasmodesmata‐ connnect protoplasts of  neighboring cells ‐ transport; relay of stimuli * symplast‐ 2 or more interconnected  l d protopolast * apoplast – cell walls, intercellular spaces  and lumen desmotubule 8
  • 9. 1/13/2010 Portions of the cell wall that remained thin   even as secondary wall is formed Primary wall only Can develop over primary pit fields f Function? TYPES: Branched simple pits (ramiform) a. simple pit Found in parenchyma cells with thickened  b. bordered pit—S‐wall develops over the pit  walls, libriform fibers, sclereids, phloem fibers cavity to form an overarching roof f h f Structure: Pit cavity / pit chamber Pit aperture Pit border Pit canal, inner and outer aperture (very thick S‐ wall) water‐conducting and mechanical xylem cells  (vessel elements, tracheids, etc.) 9
  • 10. 1/13/2010 Angiosperm Gymnosperm Pit cavity – break in S‐wall Simple pit pair Pit membrane/ closing membrane –primary  Bordered pit pair wall + middle lamella Half bordered pit pair Pit aperture Blind pit l d Unilateral compound pitting  10
  • 11. 1/13/2010 pit membrane thickening; disc shaped – porous pit membrane around the torus flexible; can go median or lateral ‐‐conifer tracheids Aspirate condition  (lateral)– latewood and all  ‐‐ occurs through  matrix dissolution heartwood Coniferales, Gnetales Round, elliptic, linear In thick cell walls:  *inner aperture becomes long and narrow *outer aperture remains circular round l d * pit canal is funnel‐shaped *fiber‐tracheid feature Scalariform Linear or crescent‐shaped thickenings  of the  Opposite primary wall and middle lamella Alternate  gymnosperms 11
  • 12. 1/13/2010 Rod shape thickenings of the wall which  Fahn, A. 1990. Plant Anatomy, 4th ed..  traverse the cell lumen radially Pergamon Press Esau, K. 1958. Plant Anatomy. John Wiley and  Sons, Inc. Evert, R. 2006. Esau’s Plant Anatomy. John  Wiley and Sons, Inc. Protopectin, pectin, pectic acid Related to pectic Plastic, amorphous colloid, hydrophilic Swelling property (hydrophilic) Hydration in young walls Impregnation starts in the intercellular  Cellulose molecules  micelles (crystal)  lamella micellar system (porous) Microfibril f consists of micellar system f And Microcapillaries liquids, lignin, waxes, cutin,  suberin, hemicellulose, pectic substances,  crystals, silica 12