PATHOLOGICAL AND REGENERATIVE PLANT ANATOMY by Professor Dr. Siegfried Fink Freiburg (Breisgau) With 1091 figures 1999 GEBRÜDER BORNTRAEGER • BERLIN • STUTTGART Contents 1. Introduction 1 2. Structural Forms of Pathological and Regenerative Changes 6 2.1 Pathological and Regenerative Changes at the Organelle Level 6 2.1.1 Plasma Membrane 8 2.1.2 Cytosol 13 2.1.3 Cytoskeleton 13 2.1.4 Nucleus 16 2.1.5 Mitochondria 22 2.1.6 Plastids 27 .7 Microbodies 42 Endoplasmic Reticulum 44 2.1.9 Ribosomes 44 2.1.10 Dictyosomes 45 2.1.11 Vacuole 46 2.1.12 Cell Wall 48 2.2 Pathological and Regenerative Changes at the Cell Level 93 2.2.1 Embryonal Cell 96 2.2.2 Mature Cell : . 96 2.2.3 Cell Malformation 96 2.2.4 Hypotrophy 97 2.2.5 Hypertrophy 99 2.2.6 Cell Division 105 2.2.7 Positional Changes 108 2.2.8 Metabolie Changes 111 2.2.8.1 Gene Activation 111 2.2.8.2 Changes in Enzyme Activities 112 2.2.8.3 Changes in Primary Metabolites 114 2.2.8.4 Changes in Secondary Metabolites 116 2.2.9 Compartmentation 124 2.2.10 Plasmolysis 135 2.2.11 Cytorrhysis 141 2.2.12 Cavitation 142 2.2.13 Cytorrhexis ... 143 2.2.14 Cytolysis 143 2.2.15 Cell Death 145 2.2.16 Regeneration 152 2.2.17 Active Exchange 154 2.2.18 Passive Withdrawal 160 2.2.19 Apoplastic Interchanges 161 2.2.20 Haustorial Interchanges 171 2.2.21 Intracellular Associations 184 2.2.22 Symplastic Interchanges 196 VIII Contents 2.2.23 Gene Transfer 202 2.2.24 Cell Fusion 213 2.3 Pathological and Regenerative Changes at the Tissue Level 215 2.3.1 Meristematic Tissue 218 2.3.2 Mature Tissue -.-.- 219 2.3.3 Target Cells . 220 2.3.4 Functional Changes 222 2.3.4.1 Dysfunction of Stomatal Regulation 222 2.3.4.2 Infiltration of Intercellular Spaces 223 2.3.4.3 Dysfunction of the Water-conducting System 224 2.3.5 Heterotopy 234 2.3.6 Directional Changes 239 2.3.7 Hypoplasia: Quantitative 244 2.3.8 Hypoplasia: Qualitative 246 2.3.9 Hyperplasia 247 2.3.10 Homeoplasia 247 2.3.11 Heteroplasia: Cataplasia 248 2.3.12 Heteroplasia: Prosoplasia 249 2.3.13 Neoplasia 250 2.3.14 Anaplasia 255 2.3.15 Metaplasia 258 2.3.16 Coenocytes 261 2.3.17 Syncytia 262 2.3.18 Intercellular Spaces 262 2.3.18.1 Non-secretory Intercellular Spaces 263 2.3.18.2 Secretory Intercellular Spaces 264 2.3.19 Separation 276 2.3.20 Compartmentätion 286 2.3.21 Regeneration 306 2.3.22 Tissue Fusion 315 2.3.23 Chimeras 327 2.3.24 Intrusion and Replacement \ 329 2.4 Pathological and Regenerative Changes at the Organ Level 332 2.4.1 Juvenile Plant 332 2.4.2 Adult Plant 335 2.4.3 Heterochrony 335 2.4.4 Correlations 336 2.4.4.1 Hormonal Correlations 336 2.4.4.2 Nutritional Correlations 337 2.4.4.3 Hydraulic Correlations 340 2.4.5 Compartmentätion 340 2.4.6 Tropisms 341 . 2.4.7 Organ Malformations 343 2.4.8 Organ Transformations 346 2.4.9 Organ Neoformations 348 2.4.10 Root Regeneration 350 2.4.10.1 Morphological Aspects 350 Contents IX 2.4.10.2 Regeneration of Damaged Root Apices 356 2.4.10.3 Hypocotyledonary and Internodal Roots 358 2.4.10.4 Nodal Roots 367 2.4.10.5 Bud-borne Roots 368 2.4.10.6 Leaf-borne Roots 370 2.4.11 Shoot Regeneration 371 2.4.11.1 Morphological Aspects 371 2A.ll.2 Regeneration of Damaged Shoot Apices 373 2.4.11.3 Induced Axillary Meristems 376 2.4.11.4 Persistent Detached Axillary Meristems 377 2.4.11.5 Suppressed Axillary Buds 379 2.4.11.6 Short Shoots 386 2.4.11.7 Long Shoots 387 2.4.11.8 Stolons 387 2.4.11.9 Rhizomes 388 2.4.11.10 Hypocotyledonary Shoots 388 2.4.11.11 Internodal Shoots 389 2.4.11.12 Root-borne Shoots (Root Suckers) 394 2.4.11.13 Leaf-borne Shoots 399 2.4.11.14 Regeneration of Leaves 401 2.4.11.15 Regeneration of Flowers 402 2.4.11.16 Vascular Connections 402 2.4.12 Embryoids 404 2.4.12.1 Morphological Aspects 404 2.4.12.2 Nucellar Embryoids 406 2.4.12.3 Suspensor-borne Embryoids 407 2.4.12.4 Shoot-borne Embryoids 407 2.4.12.5 Root-borne Embryoids 408 2.4.12.6 Leaf-borne Embryoids 408 2.4.12.7 Flower-borne Embryoids 411 2.4.12.8 Embryoids from Suspension-cultured Cells 411 3. Causes of Pathological and Regenerative Changes 423 3.1 Senescence 423 3.2 Genetic Factors 427 3.3 Physical Factors 430 3.3.1 Gravitational Stress 430 3.3.1.1 Unidirectional Exposure to Gravitation 430 3.3.1.2 Inversion 431 3.3.1.3 Centrifugation 432 3.3.1.4 Microgravity 432 3.3.1.5 Horizontal Rotation 433 3.3.2 Mechanical Stress 433 3.3.2.1 Tension 434 3.3.2.2 Pressure 434 3.3.2.3 Dynamic Loadings 435 3.3.2.4 Internal Growth Strains 436 X Contents 3.3.3 Mechanical Injuries 437 3.3.4 Chilling 456 3.3.5 Frost 457 3.3.6 Heat 472 3.3.7 Light 473 3.3.8 Ionizing Radiation 474 3.3.9 Electricity 478 3.3.10 Magnetism 479 3.3.11 Microwaves 479 3.3.12 Ultrasound 479 3.4 Chemical Factors 479 3.4.1 Water Deficiency 479 3.4.2 Water Excess 486 3.4.3 Mineral Deficiencies 489 3.4.3.1 Nitrogen Deficiency 491 3.4.3.2 Sulfur Deficiency 491 3.4.3.3 Phosphorus Deficiency 491 3.4.3.4 Calcium Deficiency 492 3.4.3.5 Potassium Deficiency 495 3.4.3.6 Magnesium Deficiency 496 3.4.3.7 Iron Deficiency 497 3.4.3.8 Manganese Deficiency 498 3.4.3.9 Zinc Deficiency 499 3.4.3.10 Copper Deficiency 499 3.4.3.11 Molybdenum Deficiency 500 3.4.3.12 Boron Deficiency 501 3.4.3.13 Multiple Deficiencies 504 3.4.4 Mineral Excesses 504 3.4.4.1 Nitrogen Excess 505 3.4.4.2 Sulfur Excess 506 3.4.4.3 Fluoride Excess 506 3.4.4.4 Aluminium Excess 506 3.4.4.5 Manganese Excess 508 3.4.4.6 Zinc Excess 509 3.4.4.7 Boron Excess 509 3.4.4.8 Copper Excess 510 3.4.4.9 Cadmium Excess 510 3.4.4.10 Cobalt Excess 511 3.4.4.11 Chromium Excess 511 3.4.4.12 Lead Excess 511 3.4.4.13 Nickel Excess 512 3.4.4.14 Uranium Excess 512 3.4.4.15 Vanadium Excess 512 3.4.4.16 NaCl Excess 513 3.4.5 Particulate Emissions 514 3.4.6 Acid Precipitation 515 3.4.7 Gaseous Air Pollutants 520 Contents XI 3.4.7.1 Sulfur Dioxide - 521 3.4.7.2 Ozone and Other Oxidants 523 3.4.7.3 Nitrogen Oxides 528 3.4.7.4 Ammonia 528 3.4.7.5 Hydrogen Fluoride 528 3.4.7.6 Hydrogen Chloride 528 3.4.7.7 Bromine 529 3.4.7.8 Hydrocarbons 529 3.4.7.9 Controlled Mixtures 529 3.4.7.10 Field Mixtures 530 3.4.8 Carbon Dioxide 531 3.4.9 Detergents 532 3.4.10 Growth Regulators and Herbicides 533 3.4.11 Fungicides 540 3.4.12 Insecticides 542 3.5 Biotic Factors 543 3.5.1 Viroids, Viruses and Phytoplasmas 543 3.5.1.1 General Aspects 543 3.5.1.2 Infection and Translocation 545 3.5.1.3 Symptoms 546 3.5.2 Bacteria 561 3.5.2.1 General Aspects 561 3.5.2.2 Infection 561 3.5.2.3 Saprotrophic Bacteria 563 3.5.2.4 Necrotrophic Bacteria 563 3.5.2.5 Biotrophic Bacteria 577 3.5.3 Fungi and Fungal-Like Organisms 589 3.5.3.1 General Aspects 589 3.5.3.2 Infection 590 3.5.3.3 Saprotrophic Fungi 598 3.5.3.4 Necrotrophic Fungi 602 3.5.3.5 Biotrophic Fungi : 628 3.5.4 Protozoa 650 3.5.5 Algae 652 3.5.6 Lichens 654 3.5.7 Parasitic Vascular Plants 655 3.5.8 Nematodes 669 3.5.9 Molluscs 678 3.5.10 Arthropods 679 3.5.11 Birds 704 3.5.12 Mammals 704 3.6 Diseases Caused by Complex or Unknown Factors 705 4. Recognition and Regulation . 720 4.1 Introduction 720 4.2 Recognition 720 4.2.1 Recognition of a Host Plant by a Parasite/Symbiont 721 XII Contents 4.2.2 Recognition of a Parasite/Symbiont by a Host Plant 722 4.2.3 Recognition of Abiotic Stress Factors 724 4.3 Signal Transduction 725 4.3.1 Signal Transduction at the Cellular Level 726 4.3.2 Long-distance Signal Transduction -.- 728 4.3.3 Signal Exchange Between Different Plants (Interplant Communi- cation) 730 4.4 Toxins 730 4.5 Hormonal Regulation 735 4.5.1 Types of Hormonal Stimuli 735 4.5.2 Changes in the Hormonal Balance by Abiotic and Biotic Factors . 737 5.