Water and Solute Permeability of Plant Cuticles: Measurement and ...

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16 1 Chemistry and Structure of Cuticles as Related to Water and Solute Permeability the cell wall. If cuticles are stained with toluidine blue at pH9, the cutin also binds toluidine blue due to the presence of carboxyl groups (unpublished results). Cationexchange capacity of cuticles will be dealt with later (Sect. 4.3). Resolution of the light microscope is of the order of 0.5µm, and this prevents the study of transversal heterogeneity of very thin cuticles. With polarised light, location and orientation of crystalline waxes have been studied. All CM investigated with polarised light exhibited birefringence or double refraction. Birefringence is evidence for the presence of crystalline structures. In cross-sections of cuticles, waxes give negative and cellulose gives a positive birefringence (Meyer 1938; Roelofsen 1952; Sitte and Rennier 1963). There are a few examples of positive birefringence due to waxes in cuticles (Sitte and Rennier 1963). Extracting or melting waxes eliminates wax birefringence. On cooling, birefringence reappears, showing re-crystallisation from the melt. Extracted CM exhibit form double refraction, indicating the presence of lamellar voids. Form birefringence disappears when the polymer matrix is imbibed with solvents having the same refractive index as cutin, which is 1.5. In periclinal positions the lamellar voids are oriented parallel to the surface of the cuticle, and in vivo they are filled with wax platelets in which the long axes of the paraffinic chains are oriented perpendicular to the surface of the CM. If viewed from the top, the waxes of Clivia cuticles appear isotropic, but near the anticlinal walls lamellae bend down towards the anticlinal walls, and in these positions waxes appear birefringent when viewed from the top (Meyer 1938). This shows an oblique orientation of the wax molecules in anticlinal pegs. The presence of cellulose in cuticles has been a subject of controversy among microscopists, because it does not exhibit the typical histochemical reactions when embedded in cutin. Crystalline cellulose exhibits positive birefringence, and the outer epidermal walls are always positive birefringent (Fig. 1.6). The CL of most cross-sections of cuticles did not show positive birefringence. Extracting or melting waxes to eliminate possible interference of negative birefringence of waxes did not change this picture. It appears that the CL of most plant cuticles contains little if any crystalline cellulose. Most researchers agree that the CP is free of crystalline cellulose. Polarised light does not detect amorphous polysaccharides, but with the transmission electron microscope (TEM) polysaccharides can be demonstrated in the CM. They are amorphous, since they are not birefringent (see below). Using Ficus elastica leaves, Sitte and Rennier (1963) observed form double refraction in the outer regions of the CM very early, when it was only about 2.5µm thick and the leaf was still unfurling. Incorporation of wax into these preformed voids occurred early, but reached its maximum only when the leaf was fully expanded. At the same time, the thickness of the CM increased by interposition of cutin between the CL and the epidermal wall, and the layered structure shown in Fig. 1.6 was formed. The mature leaf had a lamellated CP of 1–2µm, the ICL measured about 2.5µm and the ECL was about 3µm thick. The formation of voids in cutin prior to deposition of crystalline intracuticular waxes is an astounding phenomenon, yet a necessity because crystallisation of wax in a dense polymer network is improbable. Cutin monomers are likely to
1.4 Fine Structure of Cuticles 17 Fig. 1.6 Staining of cross-sections of cuticles from selected species and birefringence. The cuticle stained red with Sudan III and the cellulose wall stained blue with methylene blue. The signs of birefringence and of intensity (Γ) are shown on the left and the right side of each species respectively. (Redrawn from Sitte and Rennier 1963)
Page 2 and 3: Water and Solute Permeability of Pl
Page 4 and 5: Professor Dr. Lukas Schreiber Ecoph
Page 6 and 7: vi Preface This is not a review abo
Page 8 and 9: Contents 1 Chemistry and Structure
Page 10 and 11: Contents xi 4.6.3 Diffusion Coeffic
Page 12 and 13: Contents xiii 7.4.2 Effects of Plas
Page 14 and 15: 2 1 Chemistry and Structure of Cuti
Page 22 and 23: 10 1 Chemistry and Structure of Cut
Page 42 and 43: Chapter 2 Quantitative Description
Page 44 and 45: 2.1 Models for Analysing Mass Trans
Page 50 and 51: 2.3 Steady State Diffusion Across a
Page 52 and 53: 2.4 Steady State Diffusion of a Sol
Page 54 and 55: 2.4 Steady State Diffusion of a Sol
Page 56 and 57: 2.5 Diffusion Across a Membrane wit
Page 58 and 59: 2.5 Diffusion Across a Membrane wit
Page 60 and 61: 2.6 Determination of the Diffusion
Page 62 and 63: Solutions 51 2.7 Summary In this ch
Page 64 and 65: Chapter 3 Permeance, Diffusion and
Page 66 and 67: 3.1 Units of Permeability 55 3.1.1
Page 68 and 69: 3.1 Units of Permeability 57 identi
Page 70 and 71: 3.3 Partition Coefficients 59 The d
Page 72 and 73: Chapter 4 Water Permeability All te
Page 74 and 75: 4.1 Water Permeability of Synthetic
Page 76 and 77: 4.2 Isoelectric Points of Polymer M
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4.2 Isoelectric Points of Polymer M
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4.3 Ion Exchange Capacity 69 The hi
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4.3 Ion Exchange Capacity 71 has an
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4.3 Ion Exchange Capacity 73 Counte
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4.4 Water Vapour Sorption and Perme
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4.4 Water Vapour Sorption and Perme
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4.5 Diffusion and Viscous Transport
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4.6 Water Permeability of Isolated
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Problems 121 Our present knowledge
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Solutions 123 5. Ca 2+ , because se
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126 5 Penetration of Ionic Solutes
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146 6 Diffusion of Non-Electrolytes
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206 7 Accelerators Increase Solute
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Chapter 8 Effects of Temperature on
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8.1 Sorption from Aqueous Solutions
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8.1 Sorption from Aqueous Solutions
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8.2 Solute Mobility in Cuticles 239
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8.3 Water Permeability in CM and MX
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8.3 Water Permeability in CM and MX
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8.4 Thermal Expansion of CM, MX, Cu
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8.5 Water Permeability of Synthetic
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8.5 Water Permeability of Synthetic
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Problems 257 E D (kJ/mol) DH S (kJ/
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Chapter 9 General Methods, Sources
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9.2 Testing Integrity of Isolated C
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9.4 Distribution of Water and Solut
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9.6 Cutin and Wax Analysis and Prep
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9.7 Measuring Water Permeability 26
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9.8 Measuring Solute Permeability 2
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276 Appendix A Abbreviations (conti
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278 Appendix A Symbols (continued)
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280 Appendix B Physico-chemical pro
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Appendix C Index of Plants Botanica
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References Abraham MH, McGowan JC (
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References 287 Doty PM, Aiken WH, M
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References 289 Kolattukudy P (2004)
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References 291 Sabljic A, Güsten H
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References 293 Schreiber L, Schönh
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Index 2,4-D, 46 2,4-Dichlorophenoxy
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Index 297 leaf disks, 130 leaf surf
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Index 299 water molar volume, 110 p
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