Plant physiology and development:
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Format: | Buch |
Sprache: | English |
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New York ; Oxford
Oxford University Press
[2023]
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Ausgabe: | International seventh edition |
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Online-Zugang: | Inhaltsverzeichnis |
Beschreibung: | xxviii, 752, G-25, IC-8, I-50 Seiten Illustrationen, Diagramme |
ISBN: | 9780197614204 |
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245 | 1 | 0 | |a Plant physiology and development |c Lincoln Taiz (Professor emeritus, University of California, Santa Cruz, USA), Ian Max Møller (Associate Professor, Aarhus University, Denmark), Angus Murphy (Professor, University of Maryland), Eduardo Zeiger (Professor emeritus, University of California, Los Angeles, USA) |
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Datensatz im Suchindex
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Table of Contents UNIT i Structure and Information Systems of Plant Cells 1 CHAPTER 1 1.8 The Plant Cytoskeleton 32 The plant cytoskeleton consists of microtubules ■ Plant and Cell Architecture 3 1.1 Plant Life Processes: Unifying Principles Plant life cycles alternate between diploid and haploid generations 5 1.2 Overview of Plant Structure 7 Plant cells are surrounded by rigid cell walls 7 Plasmodesmata allow the free movement of 4 molecules between cells 7 New cells originate in dividing tissues called meristems 10 1.3 Plant Tissue Types 10 Dermal tissues cover the surfaces of plants 11 Ground tissues form the bodies of plants 12 Vascular tissues form transport networks between different parts of the plant 14 1.4 Plant Cell Compartments 15 Biological membranes are lipid bilayers that contain proteins 15 1.5 The Nucleus 18 Gene expression involves transcription, translation, and protein processing 20 Posttranslational modification of proteins determines their location, activity, and longevity 22 1.6 The Endomembrane System 23 The endoplasmic reticulum is a network of internal membranes 23 Cell wall matrix polysaccharides, secretory proteins and glycoproteins are processed in the Golgi apparatus 24 The plasma membrane has specialized regions involved in membrane recycling 26 Vacuoles have diverse functions in plant cells 27 Oil bodies are lipid-storing organelles 28 Peroxisomes play specialized metabolic roles in leaves and seeds 28 1.7 Independently Dividing Semiautonomous Organelles 29 Proplastids mature into specialized plastids in different plant tissues 30 Plastidial
and mitochondrial division are independent of nuclear division in land plants 31 and microfilaments 32 Actin, tubulin, and their polymers are in constant flux in the living cell 32 Microtubules are dynamic cylinders 34 Cytoskeletal motor proteins mediate cytoplasmic streaming and directed organelle movement 34 1.9 Cell Cycle Regulation 36 Each phase of the cell cycle has a specific set of biochemical and cellular activities 36 The cell cycle is regulated by cyclins and cyclin-dependent kinases 38 Mitosis and cytokinesis involve both microtubules and the endomembrane system 38 CHAPTER 2 ■ Cell Walls: Structure, Formation, and Expansion 45 2.1 Overview of Plant Cell Wall Functions and Structures 46 Plant cell walls vary in structure and function 46 Components differ for primary and secondary cell walls 48 Cellulose microfibrils have an ordered structure and are synthesized at the plasma membrane 50 Matrix polysaccharides are delivered to the wall via vesicles 53 Hemicelluloses are matrix polysaccharides that bind to cellulose 54 Pectins are hydrophilic gel-forming components of the primary cell wall 54 2.2 The Dynamic Primary Cell Wall 58 Primary cell walls are continually assembled during cell growth 58 2.3 Mechanisms of Cell Expansion 58 Microfibril orientation influences growth directionality of cells with diffuse growth 59 Microfibril orientation in the multilayered cell wall changes over time 60 Cortical microtubules influence the orientation of newly deposited microfibrils 60
xvi Table of Contents Many factors influence the extent and rate of cell growth 62 Stress relaxation of the cell wall drives water uptake and cell expansion 62 Leaf epidermal pavement cells provide a model for regulated cell wall expansion 63 Acid-induced growth and wall stress relaxation are mediated by expansins 63 Cell wall models are hypotheses about how molecular components fit together to make a functional wall 64 Many structural changes accompany the cessation of wall expansion 65 2.4 Secondary Cell Wall Structure and Function 66 Secondary cell walls are rich in cellulose and hemicellulose and often have a hierarchical organization 67 Lignification transforms the SCW into a hydrophobic structure resistant to deconstruction 67 CHAPTER 3 ■ Genome Structure and Gene Expression 73 3.1 Nuclear Genome Organization 73 The nuclear genome is packaged into chromatin 74 Centromeres, telomeres, and nucleolar organizer regions contain repetitive sequences 74 Transposons are mobile sequences within the genome 75 Chromosome organization is not random, in the interphase nucleus 76 Meiosis halves the number of chromosomes and allows for the recombination of alleles 76 Polyploids contain multiple copies of the entire genome 78 3.2 Plant Cytoplasmic Genomes: Mitochondria and Plastids 80 3.3 Transcriptional Regulation of Nuclear Gene Expression 81 RNA polymerase II binds to the promoter region of most protein-coding genes 81 Conserved nucleotide sequences signal transcriptional termination and polyadenylation 84 Epigenetic modifications help determine gene activity 84 3.4
Posttranscriptional Regulation of Nuclear Gene Expression 86 All RNA molecules are subject to decay 86 Noncoding RNAs regulate mRNA activity via the RNA interference (RNAi) pathway 86 3.5 Tools for Studying Gene Function 90 Mutant analysis can help elucidate gene function 90 BOX 3.1 Genetic Traits from Wild Grasses Are Used to Make Grain Crops More Resilient to Climate Change and Global Pathogen Threats 91 Molecular techniques can measure the activity of genes 92 Gene fusions can create reporter genes 92 3.6 Genetic Modification of Plants 93 3.7 Editing Plant Genomes 95 Sequence-specific nucleases induce targeted mutations 95 Gene editing can lead to precise gene replacement 97 Base editing can be used as an alternative to homology-directed repair 97 Prime editing uses an RNA repair template and reverse transcription 99 3.8 Engineering Crop Plants 99 Transgenes can confer resistance to herbicides or plant pests 99 Genetic engineering of plants remains controversial 100 CHAPTER 4 ■ Signals and Signal Transduction 103 4.1 Temporal and Spatial Aspects of Signaling 104 4.2 Signal Perception and Amplification 105 Receptors are located throughout the cell and are conserved across kingdoms 105 Signals must be amplified intracellularly to regulate their target molecules 106 Evolutionarily conserved MAP kinases amplify cellular signals 107 Evolutionarily conserved kinases regulate programmed and plastic plant development 107 Extracellular signals are perceived and transmitted by receptor-like kinases- 108 Phosphatases are the "off switch" of protein phosphorylation 109 Other protein
modifications can reconfigure cellular processes 109 Ca2+ is the most ubiquitous second messenger in plants and other eukaryotes 109 Changes in the cytosolic or cell wall pH can serve as second messengers for hormonal and stress responses 110 Reactive oxygen species act as second messengers mediat ing both environmenta l and developmental signals 111 Lipid signaling molecules act as second messengers that regulate a variety of cellular processes 111 4.3 Hormones and Plant Development 113 Auxin was discovered in early studies of coleoptile bending during phototropism 114 Gibberellins promote stem growth and were discovered in relation to the "foolish seedling disease" of rice 114 Cytokinins were discovered as cell division-promoting factors in tissue-culture experiments 116 Ethylene is a gaseous hormone that promotes fruit ripening and other developmental processes 117 Abscisic acid regulates seed maturation and stomatai closure in response to water stress 117 Brassinosteroids regulate photomorphogenesis, germina tion, and other developmental processes 118 Strigolactones suppress branching and promote rhizosphere interactions 118
Table of Contents xvii 4.4 Phytohormone Metabolism and Homeostasis 119 Indole-3-pyruvate is the primary intermediate in auxin biosynthesis 119 Gibberellins are synthesized by oxidation of the diterpene ent-kaurene 121 Cytokinins are adenine derivatives with isoprene side chains ' 123 Ethylene is synthesized from methionine via the intermediate ACC 124 Abscisic acid is synthesized from a carotenoid intermediate 125 Brassinosteroids are derived from the sterol campesterol 126 Strigolactones are synthesized from, ß-carotene 127 4.5 Movement of Hormones within the Plant 127 Plant polarity is maintained by polar auxin streams 128 Auxin transport is regulated by multiple mechanisms '131 CHAPTERS ■ Water and Plant Cells 153 5.1 Water in Plant Life 153 5.2 The Structure and Properties of Water 154 Water is a polar molecule that forms hydrogen bonds 154 Water is an excellent solvent 154 , Water has distinctive thermal properties relative to its size 155 Water has a high surface tension 155 Water has a high tensile strength 156 5.3 Diffusion and Osmosis 157 Diffusion is the net movement of molecules by random thermal agitation 157 Diffusion is most effective over short distances 158 Osmosis describes the net movement of water across a selectively permeable barrier 159 5.4 Water Potential 159 The chemical potential of water represents the free-energy status of water 159 Three major factors contribute to water potential. 159 Water potentials can be measured 160 5.5 Water Potential of Plant Cells 161 Water enters the cell along a water potential gradient 161 Water can also leave the
cell in response to a water potential gradient 162 Water potential and its components vary with growth conditions and location within the plant 163 4.6 Hormonal Signaling Pathways 132 I The cytokinin and ethylene signal transduction pathways are derived from the bacterial two-component regula tory system 132 Receptor-like kinases mediate brassinosteroid and certain auxin signaling pathways 136 The core ABA signaling components include phosphatases and kinases 136 Plant hormone signaling pathways generally employ negative regulation 139 Several plant hormone receptors include components of the ubiquitination machinery and mediate signaling via protein degradation 139 Plants have evolved mechanisms for switching off or attenuating signaling responses 143 The cellular response output to a signal is often tissue-specific 144 | Hormone responses are modulated by other endogenous molecules 144 Plants use electrical signaling for communication between tissues 146 Cross-regulation allows signal transduction pathways to be integrated 147 5.6 Cell Wall and Membrane Properties 163 Small changes in plant cell volume cause large changes in turgor pressure 163 The rate at which cells gain or lose water is influenced by plasma membrane hydraulic conductivity 164 Aquaporins facilitate the movement of water across membranes 165 5.7 Plant Water Status 166 I Physiological processes are affected by plant water status 166 Solute accumulation helps cells maintain turgor and volume 166 CHAPTER 6 . ( A W I I ■ Water Balance of Plants ■ 169 ·· 6.1 Water in the Soil 169 Soil water potential is
affected by solutes, surface tension, and gravity 170 Water moves through the soil by bulk flow 171 6.2 Water Absorption by Roots 171 Water moves in the root via the apoplast, symplasm, and transmembrane pathways 172 Solute accumulation in the xylem can generate "root pressure" 174 6.3 Water Transport through the Xylem 174 The xylem, consists of two types of transport cells 174
xviii Table of Contents Water moves through the xylem by pressure-driven bulk flow 176 Water movement through the xylem, requires a smaller pressure gradient than movement through living cells 177 What pressure difference is needed to lift water 100 meters to a treetop? 177 The cohesion-tension theory explains water transport in the xylem 178 Xylem transport of water in trees faces physical challenges 178 Plants have several mechanisms to overcome losses of xylem conductivity caused by embolism 180 6.4 Water Movement from the Leaf to the Atmosphere 181 Leaves have a large hydraulic resistance 181 The driving force for transpiration is the difference in water vapor concentration 182 Water loss is also affected by the pathway resistances 182 Stomatai control couples leaf transpiration to leaf photosynthesis 183 The cell walls of guard cells have specialized features 183 Changes in guard cell turgor pressure cause stomata to open and close 184 Internal and external signals regulate the osmotic balance of guard cells 185 The transpiration ratio measures the relationship between water loss and carbon gain 186 6.5 Overview: The Soil-Plant-Atmosphere Continuum 186 CHAPTER 7 ■ Mineral Nutrition 189 BOX 7.1 . Nitrogen Fertilizers and Climate Change CHAPTER 8 ■ Solute Transport 217 8.1 Passive and Active Transport 218 8.2 Transport of Ions across Membrane Barriers- 219 Different diffusion rates for cations and anions produce diffusion potentials 220 How does membrane potential relate to ion distribution? 220 The Nernst equation distinguishes between active and'passive transport 221
Proton transport is a major determinant of the membrane potential 222 8.3 Membrane Transport Processes 223 Channels enhance diffusion across membranes 224 Carriers bind and transport specific substances 226 Primary active transport requires energy 226 Secondary active transport is driven by ion gradients 226 Kinetic analyses can elucidate transport mechanisms 228 8,4 Membrane Transport Proteins 228 190 7.1 Essential Nutrients, Deficiencies, and Plant Disorders 191 Special techniques are used in nutritional studies 193 Nutrient solutions can sustain rapid plant growth 194 Mineral deficiencies disrupt plant metabolism and function 195 Plant tissue analysis reveals mineral deficiencies 199 BOX 7.2 Root systems differ in form but are based on common structures 205 Different areas of the root absorb mineral ions differently 207 Nutrient availability influences root growth and development 208 Mycorrhizal symbioses facilitate nutrient uptake by roots 210 Nutrients move between mycorrhizal fungi and root cells 213 lonomics: A Powerful Approach to Study Mineral Nutrition 200 7.2 Treating Nutritional Deficiencies 201 Crop yields can be improved by the addition of fertilizers 201 Some mineral nutrients can be absorbed by leaves 202 7.3 Soil, Roots, and Microbes 202 Negatively charged soil particles affect the adsorption of mineral nutrients 202 Soil pH affects nutrient availability, soil microbes, and root growth 204 Excess m ineral ions in the soil limit plant growth 204 Some plants develop extensive root systems 205 Genes encoding many transporters have been identified 230
Transporters exist for diverse nitrogen containing compounds 230 Cation transporters are diverse 231 Anion transporters have been identified 233 Transporters for metal and metalloid ions transport essential micronutrients 234 Aquaporins have diverse functions 235 Plasma membrane H+-ATPases are highly regulated P-type ATPases 236 The tonoplast H+-ATPase drives solute accumulation in vacuoles 237 H+-pyrophosphatases and P-type H+-ATPases also pump protons at the tonoplast 238 8.5 Transport in Stomatai Guard Cells 238 Blue light induces stomatal opening 239 Abscisic acid and high CO2 induce stomatal closing 240 8.6 Ion Transport in Roots 240 Solutes move through both apoplast and symplasm 240 Ions cross both symplasm and apoplast 241 Xylem parenchyma cells participate in xylem loading 242
Table of Contents CHAPTER 9 ■ Photosynthesis: The Light Reactior 9.1 Photosynthesis in Green Plants 247 9,2 General Concepts 248 Light consists of photons with characteristic energies 248 Absorption of photosynthetically active light changes the electronic states of chlorophylls 248 Photosynthetic pigments absorb the light that powers photosynthesis 250 9.3 Key Experiments in Understanding Photosynthesis 252 Action spectra relate light absorption to photosynthetic activity 252 Photosynthesis takes place in complexes containing light-harvesting antennas and photochemical reaction centers 253 The chemical reaction of photosynthesis is driven by light 254 Light drives the reduction of NADW and the formation of ATP 254 Oxygen-evolving organisms have two photosystems that operate in series 255 9.4 Organization of the Photosynthetic Apparatus 256 The chloroplast is the site of photosynthesis 256 Thylakoids contain integral membrane proteins 257 Photosystems I and II are spatially separated in the thylakoid membrane 257 Anoxygenic photosynthetic bacteria have a single reaction center 259 9.5 Organization of Light-Absorbing Antenna Systems 259 Antenna systems contain chlorophyll and are membrane-associated 259 The antenna funnels energy to the reaction center 260 Many antenna pigment-protein complexes have a common structural motif 260 9.6 Mechanisms of Electron Transport 261 Electrons from chlorophyll travel through the carriers organized in the Z scheme 261 Energy is captured when an excited chlorophyll reduces an electron acceptor molecule 263 The reaction center chlorophylls of
the two photosystems absorb at different wavelengths 264 The PSII reaction center is a multi-subunit pigmentprotein complex 264 Water is oxidized to oxygen by PSII 264 Pheophytin and two quinones accept electrons from PSII 265 Electron flow through the cytochrome b6/complex also transports protons 266 xix Plastocyanin carries electrons between the cytochrome b¿f complex and photosystem I 268 The PSI reaction center oxidizes PC and reduces ferredoxin, which transfers electrons to NADW 268 Some herbicides block photosynthetic electron flow 269 9.7 Proton Transport and ATP Synthesis in the Chloroplast 270 Cyclic electron flow augments the output of ATP to balance the chloroplast energy budget 272 9.8 Repair and Regulation of the Photosynthetic Machinery 273 Carotenoids serve as photoprotective agents 273 Some xanthophylls also participate in energy dissipation 274 The PSII reaction center is easily damaged | | I and rapidly repaired 274 Thylakoid stacking permits energy partitioning between the photosystems 275 9.9 Genetics, Assembly, and Evolution of Photosynthetic Systems 275 Chloroplast genes exhibit non-Mendelian patterns of inheritance 275 Most chloroplast proteins are imported from the cytoplasm 275 ' The biosynthesis and breakdown of chlorophyll are complex pathways 276 Complex photosynthetic organisms have evolved from, simpler forms 276 CHAPTER 10 ’ f I I I | | ' lotosynthesis: The Carbon w . Reactions 281 I 10.1 The Calvin-Benson Cycle ■ 282 The Calvin-Benson cycle has three phases: carboxylation, reduction, and regeneration 282 The fixation of CO2 via carboxylation
of ribulose 1,5-bisphosphate and the reduction of 3-phosphoglycerate yield triose phosphates 283, The regeneration of ribulose 1,5-bisphosphate ensures the continuous assimilation of CO2 284 An induction period precedes the steady state of photosynthetic CO2 assimilation 285 Many mechanisms regulate the Calvin-Benson cycle 286 Rubisco activase regulates the catalytic activity of Rubisco 287 Light regulates the Calvin-Benson cycle via the ferredoxin-thioredoxin system 288 Light-dependent ion movements modulate enzymes of the Calvin-Benson cycle 289 Light controls the assembly of chloroplast enzymes into supramolecular complexes 289
xx Table of Contents 10.2 The Oxygenation Reaction of Rubisco and Photorespiration 290 The oxygenation of ribulose 1,5-bisphosphate sets in motion photorespiration 291 Photorespiration is linked to the photosynthetic electron transport system 295 Enzymes of plant photorespiration derive from different ancestors 295 BOX 10.1 Production of Biomass May Be Enhanced by Engineering Photorespiration 296 Photorespiration interacts with many metabolic pathways 296 10.3 Inorganic Carbon-Concentrating Mechanisms 297 10.4 Inorganic Carbon-Concentrating Mechanisms: C4 Photosynthetic Carbon Fixation 297 Malate and aspartate are the primary carboxylation products of the Cį cycle 298 Kranz-type C4 plants assimilate CO2 by the concerted action of two different types of cells 299 The C4 subtypes use different mechanisms to decarbox ylate four-carbon acids transported to bundle sheath cells 301 Bundle sheath cells and mesophyll cells exhibit anatomical and biochemical differences 301 The C4 cycle also concentrates CO2 in single cells 302 Light regulates the activity of key C4 enzymes 302 Photosynthetic assimilation of CO2 in C4 plants requires more transport processes than in C3 plants 302 In hot, dry climates, the C4 cycle reduces photorespiration 303 10.5 Inorganic Carbon-Concentrating Mechanisms: Crassulacean Acid Metabolism (CAM) 303 Different mechanisms regulate C4 PEPCase and CAMPEPCase 305 CAM is a versatile mechanism sensitive to environmental stimuli 305 10.6 Accumulation and Partitioning of Photosynthates—Starch and Sucrose 305 10.7 Formation and Mobilization of Chloroplast Starch
306 Chloroplast stroma accumulates starch as insoluble granules during the day 307 Starch degradation at night requires the phosphorylation of amylopectin 310 The export of maltose prevails in the nocturnal break down of transitory starch 310 The synthesis and degradation of the starch granule are regulated by multiple mechanisms 311 10.8 Sucrose Biosynthesis and Signaling 312 Triose phosphates from the Calvin-Benson cycle build up the cy tosolic pool of three important hexose phosphates in the light 312 Fructose 2,6-bisphosphate regulates the hexose phosphate pool in the light 314 Sucrose is continuously synthesized in the cy tosol 314 Sucrose plays only a minor role in stomatai regulation 316 CHAPTER 11 lotosynthesis: Physiological and Ecological Considerations 321 11.1 Photosynthesis Is Influenced by Leaf Properties 322 Leaf anatomy and canopy structure optimize light absorption 323 Leaf angle and leaf movement can control light absorption 325 Leaves acclimate to sun and shade environments 325 11.2 Effects of Light on Photosynthesis in the Intact Leaf 326 ■ Photosynthetic light-response curves reveal differences in leaf properties 326 Leaves must dissipate excess light energy as heat 328 Absorption of too much light can lead to photoinhibition 330 11.3 . Effects of Temperature on Photosynthesis in the Intact Leaf 331 Leaves mus t dissipate vast quantities of heat 331 There is an optimal temperature for photosynthesis 332 Photosynthesis is sensitive to both high and low temperatures 332 Photosynthetic efficiency is temperature-sensitive 333 11.4 Effects of Carbon Dioxide
on Photosynthesis in the Intact Leaf 334 Atmospheric CO2 concentration keeps rising 334 CO2 diffusion to the chloroplast is essential to photosynthesis 334 CO2 supply imposes limitations on photosy nthesis 336 How will photosynthesis and respiration change in the future under elevated CO2 conditions? 338 11.5 Stable Isotopes Record Photosynthetic Properties 340 How do we measure the stable carbon isotopes of plants? 340 Why does the carbon isotope ratio vary in plants? 341 CHAPTER 12 . anslocation in the Phloem 345 12.1 Patterns of Translocation: Source to Sink 346 12.2 Pathways of Translocation 347 Sugar is translocated in phloem sieve elements 347 Mature sieve elements are living cells specialized for translocation 347 Large pores in cell walls are the prominent feature of sieve elements 348 Companion cells aid the highly specialized sieve elements 350
Table of Contents 12.3 Phloem Loading 351 Phloem loading can occur via the apoplast or symplasm 351 Apoplastic loading is characteristic of many herbaceous species 352 Sucrose loading in the apoplastic pathway requires metabolic energy 352 Phloem loading in the apoplastic pathway involves a sucrose-H+ symporter 353 Transfer cells are companion cells that are specialized for membrane transport 353 Phloem loading is symplasmic in some species 354 The oligomer-trapping model explains symplasmic loading in plants with intermediary-type companion cells 354 Phloem loading is passive in several tree species 355 The type of phloem loading is correlated with several significant characteristics 356 12.4 Long-Distance Transport: A Pressure-Driven Mechanism 357 Mass transfer is much faster than diffusion 357 The pressure-flow model is a passive mechanism for phloem transport 357 The pressure is osmotically generated 357 Some predictions of pressure: flow have been confirmed, while others require further experimentation 359 Functional sieve plate pores appear to be open channels 359 Are the pressure gradients in the sieve elements sufficient to drive phloem transport in trees? 360 Modified models for translocation by mass flow have been suggested 361 Does translocation in gymnosperms involve a different mechanism? 361 12.5 Materials Translocated in the Phloem 361 Sugars are translocated in a nonreducing form 362 , Other small organic solutes are translocated in the phloem 362 Phloem-mobile macromolecules often originate in companion cells 364 Damaged sieve elements are sealed off 364
12.6 Phloem Unloading and Sink-to-Source Transition 365 Phloem unloading and short-distance transport can occur via symplasmic or apoplastic pathways 366 Symplasmic unloading supplies growing vegetative sinks 366 Symplasmic unloading is passive but depends on energy consumption in the sink 367 Import into seeds, fruits, and storage organs often involves an apoplastic step 367 Apoplastic import is active and requires metabolic energy 368 The transition of a leaf from sink to source is gradual 369 12.7 Photosynthate Distribution: Allocation and Partitioning 371 Allocation includes storage, utilization, and transport 371 xxi Source leaves regulate allocation 371 Various sinks partition transport sugars 372 Sink tissues compete for available translocated photosynthate 372 Sink strength depends on sink size and activity 372 The source adjusts over the long term to changes in the source-to-sink ratio 373 12.8 Transport of Signaling Molecules 373 Turgor pressure and chemical signals coordinate source and sink activities 374 Mobile RNAs function as signal molecules in the phloem to regulate growth and development 374 Mobile proteins also function as signal molecules to regulate growth and development 375 Plasmodesmata function in phloem, signaling 375 BOX 12.1 Relevance of Phloem Translocation and Signaling for Climate Change and Biotechnology 376 CHAPTER 13 ՛ I I •spiration and Lipid a Metabolism 379 । m I I I . i । l Հ 13.1 Overview of Plant Respiration 379 13.2 Glycolysis 382 I । Glycolysis metabolizes carbohydrates from several sources 382 The energy-conserving phase of
glycolysis produces pyruvate, ATP, and NADH 384 Plants have alternative glycolytic reactions 385 In the absence of oxygen, fermentation regenerates the NAD+ needed for glycolytic ATP production 385 13.3 The Oxidative Pentose Phosphate Pathway 386 The oxidative pentose phosphate pathway produces NADPH and biosynthetic intermediates 386 The oxidative pentose phosphate pathway is controlled by cellular redox status 388 13.4 The Tricarboxylic Acid Cycle 388 Mitochondria are semiautonomous organelles 388 Pyruvate enters the mitochondrion and is oxidized via the TCA cycle 389 The TCA cycle of plants has unique features 391 13.5 Oxidative Phosphorylation 391 The electron transport chain catalyzes a flow of electrons from NADH to O2 392 The electron transport chain has supplementary branches 393 ATP synthesis in the mitochondrion is coupled to electron transport 394 Transporters exchange substrates and products 396 Aerobic respiration yields about 60 molecules of ATP per molecule of sucrose 396 Several subunits of respiratory complexes are encoded by the mitochondrial genome 396 Plants have several mechanisms that lower the ATP yield 398
xxii Table of Contents Respiration is an integral part of a redox and biosynthesis network 400 Respiration is controlled at multiple levels 401 Free-living and symbiotic bacteria fix nitrogen 428 Nitrogen fixation requires microanaerobic or anaerobic conditions 429 BOX 14.1 13.6 Respiration in Intact Plants and Tissues 402 Plants respire roughly half of the daily photosynthetic yield 402 Respiratory processes operate during photosynthesis 403 Different tissues and organs respire at different rates 403 BOX 13.1 Symbiotic nitrogen fixation occurs in specialized structures 430 Establishing symbiosis requires an exchange of signals 431 Nod factors produced by bacteria act as signals for symbiosis 431 Nodule formation involves phytohormones 432 The nitrogenase enzyme complex fixes N2 434 Amides and ureides are the transported forms of nitrogen 435 Modifying Respiration for Future Needs 404 Environmental factors alter respiration rates 404 13.7 Lipid Metabolism 405 Fats and oils store large amounts of energy 405 Triacylglycerols are stored in oil bodies 406 BOX 13.2 Biotechnology of Lipids in a Changing World 407 Polar glycerolipids are the main structural lipids in membranes 407 Fatty acid biosynthesis consists of cycles of two-carbon addition 407 Glycerolipids are synthesized in the plastids and the ER 410 Lipid composition influences membrane function 411 Membrane lipids are precursors of important signaling compounds 411 Storage lipids are converted into carbohydrates in germinating seeds 411 14.6 Sulfur Assimilation 435 · Sulfate is the form of sulfur transported into plants
435 Sulfate assimilation requires the reduction of sulfate to cysteine 436 Sulfate assimilation occurs mostly in leaves 438 Methionine is synthesized from cysteine 438 14.7 Phosphate Assimilation 438 miRNAs contribute to phosphate and sulfate signaling 438 14.8 Oxygen Assimilation 439 14.9 The Energetics of Nutrient Assimilation 439 CHAPTER 15 CHAPTER 14 * ssimilation of Inorganic Nutrients 417 14.1 Nitrogen in the Environment Challenges and Solutions for Solving Nitrogen Deficiency in Future Agriculture 430 418 Nitrogen passes through several forms in a biogeo chemical cycle 418 Unassimilated ammonium or nitrate I may be dangerous 419 14.2 Nitrate Assimilation 420 Many factors regulate nitrate reductase 421 Nitrite reductase converts nitrite to ammonium 421 Both roots and shoots assimilate nitrate 422 Nitrate can be transported in both xylem and phloem 422 Transceptor contributes to nitrate signaling 423 14.3 Ammonium Assimilation 424 Converting ammonium to amino acids requires two enzymes 424 Ammonium can be assimilated via an alternative pathway 426 Transamination reactions transfer nitrogen 426 Asparagine and glutamine link carbon and nitrogen metabolism 426 14.4 Amino Acid Biosynthesis 426 14.5 Biological Nitrogen Fixation 427 )iot։c Stress -՛·· ’ 15.1 Defining Plant Stress 444 Physiological adjustment to abiotic stress involves trade-offs between vegetative and reproductive development 445 15.2 Acclimation and Adaptation 445 Adaptation to stress involves genetic modification over many generations 445 Acclimation allows plants to respond to environmental fluctuations
446 15.3 Environmental Factors and Their Biological Impacts on Plants 446 Water deficit decreases turgor pressure, increases ion toxicity, and inhibits photosynthesis 447 Temperature stress affects a broad spectrum of physiological processes 447 Flooding results in anaerobic stress to the root 448 Salinity stress has both osmotic and cytotoxic effects 449 During freezing stress, extracellular ice crystal formation causes cell dehydration 449 Heavy metals can both mimic essential mineral nutrients and generate ROS 449 Ozone and ultraviolet light generate ROS that cause lesions and induce PCD 450 Combinations of abiotic stresses can induce unique signaling and metabolic pathways 450
Table of Contents xxiii Interactions occur between abiotic and biotic stresses 451 Sequential exposure to different abiotic stresses some times confers cross-protection 451 Beneficial microbes can improve plant tolerance to abiotic stress 451 15.4 Stress-Sensing Mechanisms in Plants 452 Early-acting stress sensors provide the initial signal for the stress response 452 15.5 Signaling Pathways Activated in Response to Abiotic Stress 453 The signaling intermediates of many stress-response pathways can interact 453 Acclimation to stress involves transcriptional regulatory networks called régulons 455 Chloroplasts and mitochondria respond to abiotic stress by sending stress signals to the nucleus 456 Plant-wide waves of Ca2+ and ROS mediate systemic acquired acclimation 456 Epigenetic mechanisms, retrotransposons, and small RNAs provide additional protection against stress 456 Hormonal interactions regulate abiotic stress responses 459 CHAPTER 16' ■ Signals from Sunlight 475 16.1 Plant Photoreceptors 476 ՛ Photoresponses are driven by light quality or spectral properties of the energy absorbed 477 Plants responses to light can be distinguished by the amount of light required 478 16.2 Phytochromes 480 Phytochrome is the primary photoreceptor for red and far-red light 480 Phytochrome can interconvert between Pr and Pfr forms 480 Pfr is the physiologically active form of phytochrome 481 The phytochrome chromophore and protein both undergo conformational changes in response to red light 481 Pfr is partitioned between the cytosol and the nucleus 483 16.3 Phytochrome Responses 484
Phytochrome responses vary in lag time and escape time 484 Phytochrome responses fall into three main categories based on the amount of light required 484 Phytochrome A mediates responses to continuous far-red light 486 15.6 Physiological and Developmental A Mechanisms That Protect Plants against Abiotic Stress 460 Plants adjust osmotically to drying soils by accumulating solutes 460 Submerged organs develop aerenchyma tissue in response to hypoxia 461 Antioxidants and ROS-scavenging pathways protect cells from oxidative stress 462 Molecular chaperones and molecular shields protect proteins and membranes during abiotic stress 462 Plants can alter their membrane lipids in response to temperature and other abiotic stresses 463 Exclusion and internal tolerance mechanisms allow plants to cope with toxic ions 464 Phytochelatins and other chelators contribute to internal tolerance of toxic metal ions 465 Plants use cryoprotectant molecules and antifreeze proteins to prevent ice crystal formation 466 ABA signaling during water stress causes the massive efflux of K+ and anions from guard cells 466 Plants can alter their morphology in response to abiotic stress 467 The process of recovery from stress can be dangerous to the plant and requires a coordinated adjustment of plant metabolism and physiology 469 Phytochrome В mediates responses to continuous red or white light 486 Roles for phytochromes C, D, and E are emerging 486 16.4 Phytochrome Signaling Pathways 487 Phytochrome regulates membrane potentials and ion fluxes 487 Phy tochrome regulates gene expression 487 Phytochrome
interacting factors (PIFs) act early in signaling 488 ' . Phytochrome signaling involves protein phosphorylation and déphosphorylation 488 Phytochrome-induced photomorphogenesis involves pro tein degradation 489 16.5 Blue-Light Responses and Photoreceptors 490 . A Blue-light responses have characteristic kinetics and lag times 490 16.6 Cryptochromes 491 The activated FAD chromophore of cryptochrome causes a conformational change in the protein 491 cryl and cry2 have different developmental effects 492 Nuclear cryptochromes inhibit COPl-induced protein degradation 493 Cryptochrome can also bind to transcriptional regulators directly 493
xxiv Table of Contents 16.7 Interactions of Cryptochrome with Other Photoreceptors 493 Stem elongation is inhibited by both red and blue photoreceptors 493 Phytochrome interacts with cryptochrome to regulate flowering 494 The circadian clock is regulated by multiple aspects of light 494 16.8 Phototropins 495 Blue light induces changes in FMN absorption maxima associated with conformation changes 495 The LOV2 domain is primarily responsible for kinase activation in response to blue light 496 Blue light induces a conformational change that "uncages" the kinase domain of phototropin and leads to autophosphorylation 496 Phototropins trigger plant movements that enhance light use 496 Blue light initiates stomatai opening via activation of the plasma membrane H+-ATPase 498 16.9 Responses to Ultraviolet Radiation 500 CHAPTER 17 ■ Seed Dormancy, Germination, and Seedling Establishment 505 17.1 Seed Structure 506 Seed anatomy varies widely among different plant groups 506 17.2 Seed Dormancy 508 There are two basic types of seed dormancy mechanisms: exogenous and endogenous 508 Non-dormant seeds can exhibit vivipary and precocious germination 509 The ABA:GA ratio is the primary determinant of embry onic seed dormancy 510 17.3 Release from Dormancy 511 Light is an important signal that breaks dormancy in small seeds 511 Some seeds require either chilling or after-ripening to break dormancy 511 Seed dormancy can be broken by various chemical compounds 512 17.4 Seed Germination 512 Germination and postgermination can be divided into three phases corresponding to the phases of water
uptake 513 17.5 Mobilization of Stored Reserves 514 Cereal seeds are a model for understanding starch mobilization 515 Legume seeds are a model for understanding protein mobilization 516 Oilseeds are a model for understanding lipid remobilization 517 17.6 Seedling Growth and Establishment 517 The development of emerging seedlings is strongly influenced by light 517 Gibberellins and brassinosteroids both suppress photomorphogenesis in darkness 518 Hook opening is regulated by phytochrome, auxin, and ethylene 519 Vascular differentiation begins during seedling emergence 520 The root tip has specialized cells 520 Ethylene and other hormones regulate root hair development 521 17.7 Differential Growth Enables Successful Seedling Establishment 522 Ethylene affects microtubule orientation and induces lateral cell expansion 523 Auxin promotes growth in stems and coleoptiles, while inhibiting growth in roots 524 The minimum lag time for auxin-induced elongation is 10 minutes 525 Auxin-induced proton extrusion loosens the cell wall 526 17.8 Tropisms: Growth in Response to Directional Stimuli 526 Gravitropism involves the lateral redistribution •af auxin 526 The gravitropic stimulus perturbs the symmetric move ments of auxin 526 Gravity perception is triggered by the sedimentation of amyloplasts 529 Gravity sensing may involve pH and calcium ions (Ca2+) as second messengers 532 Thigmotropism involves signaling by Ca2+, pH, and reac tive oxygen species 533 Hydrotropism involves ABA signaling and asymmetric cytokinin responses 534 Phototropins are the light receptors involved in
phototropism 535 Phototropism is mediated by the lateral redistribution of auxin 535 Shoot phototropism occurs in a series of steps 536 CHAPTER 18 ■ Vegetative Growth and Organogenesis: Primary Growth of the Plant Axis 541 18.1 Meristematic Tissues: Foundations for Indeterminate Growth 541 The root and shoot apical meristems use similar strategies to enable indeterminate growth 542 18.2 The Root Apical Meristem 542 The root tip has four developmental zones 542 The origin of different root tissues can be traced to specific initial cells 543 Auxin and cytokinin contribute to the maintenance and function of the RAM 543 18.3 The Shoot Apical Meristem 545 The shoot apical meristem has distinct zones and layers 545 A combination of positive and negative interactions deter mines apical meristem size 546
xxv Table of Contents KNOX class homeodomain transcription factors help maintain proliferation in the SAM through regulation of cytokinin and GA concentrations 547 Localized auxin accumulation promotes leaf initiation 547 Axillary meristems form in the axils of leaf primordia 548 18.4 Leaf Development 549 Growth determines leaf shape 551 18.5 The Establishment of Leaf Polarity 551 A signal from the SAM initiates adaxial-abaxial polarity 551 Antagonism between sets of transcription factors determines adaxial-abaxial leaf polarity 552 MYB transcription factors, HD-ZIP HI proteins, and KN0X1 repression promote adaxial identity 552 Abaxial identity is determined by auxin, KANADI, and YABBY 552 Blade outgrowth is auxin dependent and regulated by the YABBY and WOX genes 553 Leaf proximal-distal polarity also depends on specific gene expression 553 . In compound leaves, de-repression of the KNOX1 gene promotes leaflet formation 554 18.6 Differentiation of Epidermal Cell Types 555 Guard cell identity is determined by a specialized epidermal lineage 555 Two groups of bHLH transcription factors govern stomata! cell identity transitions 556 Cell-to-cell peptide signals regulate stornata! patterning 557 Intrinsic polarity in the stomatai lineage aids stornata! spacing 557 Environmental factors also regulate stomata! density 558 Stomata development in monocots involves some genes that are orthologous to those in Arabidopsis 558 18.7 Venation Patterns in Leaves 559 The primary leaf vein is initiated discontinuously from the preexisting vascular system 560 Auxin canalization initiates
development of the leaf trace 560 Basipetal auxin transport from the LI layer of the leaf primordium initiates development of the leaf trace procambium 561 The existing vasculature guides the growth of the leaf trace 562 Vascular development proceeds from procambium differentiation 562 Higher-order leaf veins differentiate in a predictable hierarchical order 562 Auxin regulates higher-order vein formation and patterning 563 CHAPTER 19 I I ■ Vegetative Growth and Organogenesis: Branching and Secondary Growth 567 19.1 Shoot Branching and Architecture 568 Auxin, cytokinins, and strigolactones regulate axillary bud outgrowth 569 Auxin from the shoot tip maintains | apical dominance 569 Strigolactones act locally to repress axillary bud growth 571 I Cytokinins antagonize the effects of strigolactones 571 Integration of environmental and hormonal branching signals is required for plant fitness 572 Axillary bud dormancy is affected by season, position, and age factors 572 19.2 Root Branching and Architecture 573 Lateral root primordia arise from the xylem pole pericycle cells 573 Lateral root formation can be divided into four distinct stages 574 Lateral root founder cells undergo asymmetric cell divi sions to initiate formation of lateral root primordia 576 Monocots and eudicots differ in their predominant root types 576 Transcription factors regulate the gravitropic setpoint angles of lateral roots and shoots 577 Plants can modify their root system architecture to optimize water and nutrient uptake 578 19,3 Secondary Growth 578 Two types of lateral meristems are involved in
secondary growth 578 The vascular cambium produces secondary xylem and phloem 579 Mobile transcription factors pre-pattern the vascular cambium 580 ' The gene networks that control secondary meristems share similarities and differences with those that control the apical meristems 582 Several phytohormones regulate vascular cambium activity and differentiation of secondary xylem and phloem 584 The cork cambium gives rise to the outer corky layer called the periderm 585 Bark has diverse protective and storage functions 586 Epicormic buds covered by bark can sprout after forest fires 586 CHAPTER 20 ' I I | ■ The Control of Flowering and Floral Development 591 Հ I ' : 20.1 Floral Evocation: Integrating ■ Environmental Cues 591 ' 20.2 The Shoot Apex and Phase Changes 592 Plants progress through three developmental phases 592 Juvenile tissues are produced first and are located at the base of the shoot 592 Phase changes can be influenced by nutrients, gibberel lins, and other signals 593 20.3 Circadian Rhythms: The Clock Within 594 Circadian rhythms exhibit characteristic features 595 Phase shifting adjusts circadian rhythms to different day-night cycles 597 Phytochromes and cryptochromes entrain the clock 597
xxvi Table of Contents 20.4 Photoperiodism: Monitoring Day Length 597 Plants can be classified according to their photoperiodic responses 597 The leaf is the site of perception of the photoperiodic signal 599 The length of the night is important for floral induction 599 Night breaks can cancel the effec t of the dark period 600 Photoperiodic timekeeping during the night depends on a circadian clock 601 The external coincidence model is based on oscillating light sensitivity 602 The coincidence of CONSTANS expression and light promotes flowering in LDPs 602 SDPs use a coincidence mechanism to inhibit flowering in long days 603 BOX 20.1 Refining Molecular Mechanisms of Photoperiodic Flowering Happening in Natural Environments 604 Phytochrome is the primary photoreceptor in photoperiodism 605 A blue-light photoreceptor regulates flowering in some LDPs 606 20.5 Long-Distance Signaling Involved in Flowering 606 Grafting studies provided the first evidence for a. trans missible floral stimulus 607 Florigen is translocated in the phloem 608 20.6 The Identification of Florigen 608 The Arabidopsis protein FLOWERING LOCUS T (FT) is florigen 608 20.7 Vernalization: Promoting Flowering with Cold 610 Vernalization results in competence to flower at the shoot apical meristem 610 Vernalization can involve epigenetic changes in gene expression 611 A range of vernalization pathways may have evolved. 612 20.8 Multiple Pathways Involved in Flowering 612 Gibberellins and ethylene can induce flowering 612 The transition to flowering involves multiple factors and pathways 613 20.9 Floral
Meristems and Floral Organ Development 613 The shoot apical meristem in Arabidopsis changes with development 613 The four different types of floral organs are initiated as separate whorls 614 ' Two major categories of genes regulate floral development 615 Floral meristem identity genes regulate meristem function . 615 Homeotic mutations led to the identification of floral organ identity genes 616 The ABC model partially explains the determination of . floral organ identity 617 Arabidopsis Class E genes are required for the activities of the A, B, and C genes 618 According to the Quartet Model, floral organ identity is regulated by tetrameric complexes of the ABCE proteins 619 Class D genes are required for ovule formation 620 Floral asymmetry in flowers is regulated by gene expression 620 CHAPTER 21 ’ ’ ■ Sexual Reproduction: From Gametes to Fruits 625 21.1 Development of the Male and Female Gametophyte Generations 625 21.2 Formation of Male Gametophytes ' in the Stamen 627 Pollen grain formation occurs in two successive stages 627 The multilayered pollen cell wall is surprisingly complex 629 21.3 Female Gametophyte Development in the Ovule 630 The Arabidopsis gynoecium is an important model system for studying ovule development 631 The vast majority of angiosperms exhibit Polygonum-type embryo sac development 632 Functional megaspores undergo a series of free nuclear mitotic divisions followed by cellularization 632 21.4 Pollination and Fertilization in Flowering Plants 633 The progamic phase includes everything from pollen landing and tube growth to the fusion of sperm
and egg 633 Adhesion and hydration of a pollen grain on a compati ble flower depend on recognition between pollen and stigma surfaces 634 Ca2+֊triggered polarization of the pollen grain precedes tube formation 635 Pollen tubes grow by tip growth 636 Receptor-like kinases are thought to regulate the ROPI GTPase switch, a master regulator of tip growth 638 Pollen tube tip growth in the pistil is guided by both physical and chemical cues 638 Style tissue may condition pollen tubes to grow toward the embryo sac 639 Synergid cells release chemoattractants that guide pollen tube growth to the micropyle 640 Double fertilization occurs in three distinct stages 641 21.5 Selfing versus Outcrossing 642 Hermaphroditic and monoecious species have evolved floral features to ensure outcrossing 642 Cytoplasmic male sterility (CMS) occurs in the wild and is of great utility in agriculture 642 Self-incompatibility (SI) is the primary mechanism that enforces outcrossing in angiosperms 643
Table of Contents xxvii Two distinct genetic mechanisms govern self-incompatibility 644 The Brassicaceae sporophytic SI system is mediated by S locus-encoded receptors and ligands 645 Cytotoxic S-RNases and F-box proteins determine gametophytic self-incompatibility (GSI) 645 21.6 Apomixis: Asexual Reproduction by Seed . 647 Apomixis is not an evolutionary dead end 647 21.7 Endosperm Development 647 Cellularization of coenocytic endosperm in Arabidopsis progresses from the micropylar to the chalazal region 648 Cellularization of the coenocytic endosperm of cereals progresses centripetally 649 Endosperm development and embryogenesis can occur autonomously 650 Many of the genes that control endosperm development are differentially expressed maternal or paternal genes 651 Cells of the starchy endosperm and aleurone layer follow divergent developmental pathways 652 21.8 Seed Coat Development 652 Seed coat development appears to be regulated by the endosperm 652 21.9 Seed Maturation and Desiccation Tolerance 653 Seed filling and desiccation tolerance phases overlap in most species 654 The acquisition of desiccation tolerance involves many metabolic pathways 654 During the acquisition of desiccation tolerance, the cells of the embryo acquire aiglassy state 655 LEA proteins and nonreducing sugars have been impli cated in seed desiccation tolerance 655 Abscisic acid plays a key role in seed maturation 655 Coat-imposed dormancy is correlated with long-term ' seed viability 655 21.10 Fruit Development and Ripening 656 The phytohormones auxin and gibberellic acid (GA) regulate fruit
set and parthenocarpy 656 Specific transcription factors regulate the development of the dehiscence zone 658 Tomato is an important model system for studying fleshy fruit development 659 Fleshy fruits undergo ripening 660 Ripening involves changes in the color of fruit 660 Fruit softening involves the coordinated action of many cell wall-degrading enzymes 661 Taste and flavor reflect changes in acids, sugars, aroma, and other compounds 661 The causal link between ethylene and ripening was demonstrated in transgenic and mutant tomatoes 662 Climacteric and non-climacteric fruit differ in their ethylene responses 662 The ripening process is transcriptionally regulated 663 Studying the molecular mechanism of ripening can have commercial applications 664 CHAPTER 22 I I I I ■ Embryogenesis: The Origin of Plant Architecture 669 22.1 Embryogenesis in Monocots I I and Eudicots 670 I I . Embryogenesis differs between monocots and eudicots, but also shares common features 670 22.2 Establishment of Apical-Basal Polarity 672 Apical-basal polarity is established early in embryogenesis 672 Zygote polarization can be studied using live imaging 673 22.3 Mechanisms Guiding Embryogenesis 676 Intercellular signaling processes play key roles in guiding position-dependent development 677 Cell-cell communication during early embryo develop ment may be regulated by plasmodesmata 677 Mutant analyses have identified genes for signaling pro cesses that are essential for embryo organization 678 22.4 Auxin Signaling During Embryogenesis 680 Spatial patterns of auxin accumulation regulate key
developmental events 680 The GNOM protein establishes a polar distribution of PIN auxin efflux proteins 681 MONOPTEROS encodes a transcription factor that is activated by auxin 681 22,5 Radial Patterning During Embryogenesis 682 I | Procambial precursors for the vascular stele lie at the center of the radial axis 683 The differentiation of cortical and endodermal cells involves the intercellular movement of a transcription factor 684 22,6 Formation of the Root and Shoot Apical Meristems 686 Root formation involves MONOPTEROS and other auxin-regulated transcription factors 686 Shoot formation requires HD-ZIP III, SHOOT MERISTEMLESS, and WUSCHEL genes 687 Plants can initiate embryogenesis in multiple types of cells 687 CHAPTER 23 I I I I | ■ Plant Senescence and Developmental Cell Death '691 23.1 Programmed Cell Death 692 I I Distinct types of PCD occur in plants 693 Developmental PCD and pathogen-triggered PCD involve distinct processes 693 The autophagy pathway captures and degrades cellular constituents within lytic compartments 693 Autophagy plays a dual role in the regulation of plant PCD 695 Autophagy is required for nutrient recycling during plant senescence 696
xxviii Table of Contents 23.2 The Leaf Senescence Syndrome 696 Leaf senescence may be sequential, seasonal, or stress-induced 697 Leaves undergo massive structural and biochemical changes during leaf senescence 698 The autolysis of chloroplast proteins occurs in multiple compartments 698 The STAY-GREEN (SGR) protein is required for both LHCP II protein recycling and chlorophyll catabolism 699 23.3 Regulation of Leaf Senescence: A Multi-Layered Network 700 Leaf senescence depends on the comprehensive regulation of pathways that respond to endogenous and environ mental factors 701 Plant hormones and other signaling agents can act as positive or negative regulators of leaf senescence 706 Positive senescence regulators 707 Negative senescence regulators 708 23.4 Abscission 709 Organ abscission is regulated by developmental and environmental cues 711 23.5 Whole-Plant Senescence 713 Angiosperm life cycles may be annual, biennial, or perennial 714 Whole-plant senescence differs from aging in animals 714 The determinacy of shoot apical meristems is develop mentally regulated 715 Nutrient redistribution may trigger senescence in monocarpic plants 716 The productivity of tall trees continues to increase right up to the onset of senescence 716 CHAPTER 24 ■ Biotic Interactions 721 24.1 Plant Interactions with Beneficial Microorganisms 723 Nod factors are recognized by the Nod factor receptor (NFR) in legumes 723 Arbuscular mycorrhizal associations and nitrogen-fixing symbioses involve related signaling pathways 723 Rhizobacteria can increase nutrient availability, stimulate root
branching, and protect against pathogens 725 24.2 Herbivore Interactions That Harm Plants 725 Mechanical barriers provide a first line of defense against insect pests and pathogens 726 Plant specialized metabolites can deter insect herbivores 728 Plants store constitutive toxic compounds in specialized structures 728 Plants often store defense chemicals as nontoxic watersoluble sugar conjugates in the vacuole 730 24.3 Inducible Defense Responses to Insect . Herbivores 732 Plants can recognize specific components of insect saliva 733 Ca2+ signaling and activation of the MAP kinase pathway are early events associated with insect herbivory 734 Jasmonate activates defense responses against insect herbivores 734 Jasmonate acts through a conserved ubiquitin ligase signaling mechanism 735 Hormonal interactions contribute to planfoinsect herbi vore interactions 735 JA initiates the production of defense proteins that inhibit herbivore digestion 736 Herbivore damage induces systemic defenses 736 Glutamate receptor-like (GLR) genes are required for long-distance electrical signaling during herbivory 737 Herbivore-induced volatiles can repel herbivores and attract natural enemies 738 Herbivore-induced volatiles can serve as long-distance signals between plants 739 Herbivore-induced volatiles can also act as systemic ., signals within a plant 739 Defense responses to herbivores and pathogens are regulated by circadian rhythms 739 Insects have evolved mechanisms to defeat plant defenses 741 24.4 Plant Defenses against Pathogens 741 Microbial pathogens have evolved various strategies to
invade host plants 741 Pathogens produce effector molecules that aid in the colonization of their plant host cells 742 Plants can detect pathogens through perception of pathogen-derived "danger signals" 743 R genes provide resistance to individual pathogens by recognizing strain-specific effectors 744 The hypersensitive response is a common defense against pathogens 745 A single encounter with a pathogen may increase resistance to future attacks 746 The main components of the salicylic acid signaling pathway have been identified 746 Phytoalexins with antimicrobial activity accumulate after pathogen attack 747 RNA interference plays a central role in antiviral immune responses in plants 747 Some plant parasitic nematodes form specific associations through the formation of distinct feeding structures 748 Plants compete with other plants by secreting allelopathic specialized metabolites into the soil 749 Some plants are parasites of other plants 749 Glossary G-1 Illustration Credits IC-1 Index 1-1 |
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Table of Contents UNIT i Structure and Information Systems of Plant Cells 1 CHAPTER 1 1.8 The Plant Cytoskeleton 32 The plant cytoskeleton consists of microtubules ■ Plant and Cell Architecture 3 1.1 Plant Life Processes: Unifying Principles Plant life cycles alternate between diploid and haploid generations 5 1.2 Overview of Plant Structure 7 Plant cells are surrounded by rigid cell walls 7 Plasmodesmata allow the free movement of 4 molecules between cells 7 New cells originate in dividing tissues called meristems 10 1.3 Plant Tissue Types 10 Dermal tissues cover the surfaces of plants 11 Ground tissues form the bodies of plants 12 Vascular tissues form transport networks between different parts of the plant 14 1.4 Plant Cell Compartments 15 Biological membranes are lipid bilayers that contain proteins 15 1.5 The Nucleus 18 Gene expression involves transcription, translation, and protein processing 20 Posttranslational modification of proteins determines their location, activity, and longevity 22 1.6 The Endomembrane System 23 The endoplasmic reticulum is a network of internal membranes 23 Cell wall matrix polysaccharides, secretory proteins and glycoproteins are processed in the Golgi apparatus 24 The plasma membrane has specialized regions involved in membrane recycling 26 Vacuoles have diverse functions in plant cells 27 Oil bodies are lipid-storing organelles 28 Peroxisomes play specialized metabolic roles in leaves and seeds 28 1.7 Independently Dividing Semiautonomous Organelles 29 Proplastids mature into specialized plastids in different plant tissues 30 Plastidial
and mitochondrial division are independent of nuclear division in land plants 31 and microfilaments 32 Actin, tubulin, and their polymers are in constant flux in the living cell 32 Microtubules are dynamic cylinders 34 Cytoskeletal motor proteins mediate cytoplasmic streaming and directed organelle movement 34 1.9 Cell Cycle Regulation 36 Each phase of the cell cycle has a specific set of biochemical and cellular activities 36 The cell cycle is regulated by cyclins and cyclin-dependent kinases 38 Mitosis and cytokinesis involve both microtubules and the endomembrane system 38 CHAPTER 2 ■ Cell Walls: Structure, Formation, and Expansion 45 2.1 Overview of Plant Cell Wall Functions and Structures 46 Plant cell walls vary in structure and function 46 Components differ for primary and secondary cell walls 48 Cellulose microfibrils have an ordered structure and are synthesized at the plasma membrane 50 Matrix polysaccharides are delivered to the wall via vesicles 53 Hemicelluloses are matrix polysaccharides that bind to cellulose 54 Pectins are hydrophilic gel-forming components of the primary cell wall 54 2.2 The Dynamic Primary Cell Wall 58 Primary cell walls are continually assembled during cell growth 58 2.3 Mechanisms of Cell Expansion 58 Microfibril orientation influences growth directionality of cells with diffuse growth 59 Microfibril orientation in the multilayered cell wall changes over time 60 Cortical microtubules influence the orientation of newly deposited microfibrils 60
xvi Table of Contents Many factors influence the extent and rate of cell growth 62 Stress relaxation of the cell wall drives water uptake and cell expansion 62 Leaf epidermal pavement cells provide a model for regulated cell wall expansion 63 Acid-induced growth and wall stress relaxation are mediated by expansins 63 Cell wall models are hypotheses about how molecular components fit together to make a functional wall 64 Many structural changes accompany the cessation of wall expansion 65 2.4 Secondary Cell Wall Structure and Function 66 Secondary cell walls are rich in cellulose and hemicellulose and often have a hierarchical organization 67 Lignification transforms the SCW into a hydrophobic structure resistant to deconstruction 67 CHAPTER 3 ■ Genome Structure and Gene Expression 73 3.1 Nuclear Genome Organization 73 The nuclear genome is packaged into chromatin 74 Centromeres, telomeres, and nucleolar organizer regions contain repetitive sequences 74 Transposons are mobile sequences within the genome 75 Chromosome organization is not random, in the interphase nucleus 76 Meiosis halves the number of chromosomes and allows for the recombination of alleles 76 Polyploids contain multiple copies of the entire genome 78 3.2 Plant Cytoplasmic Genomes: Mitochondria and Plastids 80 3.3 Transcriptional Regulation of Nuclear Gene Expression 81 RNA polymerase II binds to the promoter region of most protein-coding genes 81 Conserved nucleotide sequences signal transcriptional termination and polyadenylation 84 Epigenetic modifications help determine gene activity 84 3.4
Posttranscriptional Regulation of Nuclear Gene Expression 86 All RNA molecules are subject to decay 86 Noncoding RNAs regulate mRNA activity via the RNA interference (RNAi) pathway 86 3.5 Tools for Studying Gene Function 90 Mutant analysis can help elucidate gene function 90 BOX 3.1 Genetic Traits from Wild Grasses Are Used to Make Grain Crops More Resilient to Climate Change and Global Pathogen Threats 91 Molecular techniques can measure the activity of genes 92 Gene fusions can create reporter genes 92 3.6 Genetic Modification of Plants 93 3.7 Editing Plant Genomes 95 Sequence-specific nucleases induce targeted mutations 95 Gene editing can lead to precise gene replacement 97 Base editing can be used as an alternative to homology-directed repair 97 Prime editing uses an RNA repair template and reverse transcription 99 3.8 Engineering Crop Plants 99 Transgenes can confer resistance to herbicides or plant pests 99 Genetic engineering of plants remains controversial 100 CHAPTER 4 ■ Signals and Signal Transduction 103 4.1 Temporal and Spatial Aspects of Signaling 104 4.2 Signal Perception and Amplification 105 Receptors are located throughout the cell and are conserved across kingdoms 105 Signals must be amplified intracellularly to regulate their target molecules 106 Evolutionarily conserved MAP kinases amplify cellular signals 107 Evolutionarily conserved kinases regulate programmed and plastic plant development 107 Extracellular signals are perceived and transmitted by receptor-like kinases- 108 Phosphatases are the "off switch" of protein phosphorylation 109 Other protein
modifications can reconfigure cellular processes 109 Ca2+ is the most ubiquitous second messenger in plants and other eukaryotes 109 Changes in the cytosolic or cell wall pH can serve as second messengers for hormonal and stress responses 110 Reactive oxygen species act as second messengers mediat ing both environmenta l and developmental signals 111 Lipid signaling molecules act as second messengers that regulate a variety of cellular processes 111 4.3 Hormones and Plant Development 113 Auxin was discovered in early studies of coleoptile bending during phototropism 114 Gibberellins promote stem growth and were discovered in relation to the "foolish seedling disease" of rice 114 Cytokinins were discovered as cell division-promoting factors in tissue-culture experiments 116 Ethylene is a gaseous hormone that promotes fruit ripening and other developmental processes 117 Abscisic acid regulates seed maturation and stomatai closure in response to water stress 117 Brassinosteroids regulate photomorphogenesis, germina tion, and other developmental processes 118 Strigolactones suppress branching and promote rhizosphere interactions 118
Table of Contents xvii 4.4 Phytohormone Metabolism and Homeostasis 119 Indole-3-pyruvate is the primary intermediate in auxin biosynthesis 119 Gibberellins are synthesized by oxidation of the diterpene ent-kaurene 121 Cytokinins are adenine derivatives with isoprene side chains ' 123 Ethylene is synthesized from methionine via the intermediate ACC 124 Abscisic acid is synthesized from a carotenoid intermediate 125 Brassinosteroids are derived from the sterol campesterol 126 Strigolactones are synthesized from, ß-carotene 127 4.5 Movement of Hormones within the Plant 127 Plant polarity is maintained by polar auxin streams 128 Auxin transport is regulated by multiple mechanisms '131 CHAPTERS ■ Water and Plant Cells 153 5.1 Water in Plant Life 153 5.2 The Structure and Properties of Water 154 Water is a polar molecule that forms hydrogen bonds 154 Water is an excellent solvent 154 , Water has distinctive thermal properties relative to its size 155 Water has a high surface tension 155 Water has a high tensile strength 156 5.3 Diffusion and Osmosis 157 Diffusion is the net movement of molecules by random thermal agitation 157 Diffusion is most effective over short distances 158 Osmosis describes the net movement of water across a selectively permeable barrier 159 5.4 Water Potential 159 The chemical potential of water represents the free-energy status of water 159 Three major factors contribute to water potential. 159 Water potentials can be measured 160 5.5 Water Potential of Plant Cells 161 Water enters the cell along a water potential gradient 161 Water can also leave the
cell in response to a water potential gradient 162 Water potential and its components vary with growth conditions and location within the plant 163 4.6 Hormonal Signaling Pathways 132 I The cytokinin and ethylene signal transduction pathways are derived from the bacterial two-component regula tory system 132 Receptor-like kinases mediate brassinosteroid and certain auxin signaling pathways 136 The core ABA signaling components include phosphatases and kinases 136 Plant hormone signaling pathways generally employ negative regulation 139 Several plant hormone receptors include components of the ubiquitination machinery and mediate signaling via protein degradation 139 Plants have evolved mechanisms for switching off or attenuating signaling responses 143 The cellular response output to a signal is often tissue-specific 144 | Hormone responses are modulated by other endogenous molecules 144 Plants use electrical signaling for communication between tissues 146 Cross-regulation allows signal transduction pathways to be integrated 147 5.6 Cell Wall and Membrane Properties 163 Small changes in plant cell volume cause large changes in turgor pressure 163 The rate at which cells gain or lose water is influenced by plasma membrane hydraulic conductivity 164 Aquaporins facilitate the movement of water across membranes 165 5.7 Plant Water Status 166 I Physiological processes are affected by plant water status 166 Solute accumulation helps cells maintain turgor and volume 166 CHAPTER 6 . ( A W I I ■ Water Balance of Plants ■ 169 ·· 6.1 Water in the Soil 169 Soil water potential is
affected by solutes, surface tension, and gravity 170 Water moves through the soil by bulk flow 171 6.2 Water Absorption by Roots 171 Water moves in the root via the apoplast, symplasm, and transmembrane pathways 172 Solute accumulation in the xylem can generate "root pressure" 174 6.3 Water Transport through the Xylem 174 The xylem, consists of two types of transport cells 174
xviii Table of Contents Water moves through the xylem by pressure-driven bulk flow 176 Water movement through the xylem, requires a smaller pressure gradient than movement through living cells 177 What pressure difference is needed to lift water 100 meters to a treetop? 177 The cohesion-tension theory explains water transport in the xylem 178 Xylem transport of water in trees faces physical challenges 178 Plants have several mechanisms to overcome losses of xylem conductivity caused by embolism 180 6.4 Water Movement from the Leaf to the Atmosphere 181 Leaves have a large hydraulic resistance 181 The driving force for transpiration is the difference in water vapor concentration 182 Water loss is also affected by the pathway resistances 182 Stomatai control couples leaf transpiration to leaf photosynthesis 183 The cell walls of guard cells have specialized features 183 Changes in guard cell turgor pressure cause stomata to open and close 184 Internal and external signals regulate the osmotic balance of guard cells 185 The transpiration ratio measures the relationship between water loss and carbon gain 186 6.5 Overview: The Soil-Plant-Atmosphere Continuum 186 CHAPTER 7 ■ Mineral Nutrition 189 BOX 7.1 . Nitrogen Fertilizers and Climate Change CHAPTER 8 ■ Solute Transport 217 8.1 Passive and Active Transport 218 8.2 Transport of Ions across Membrane Barriers- 219 Different diffusion rates for cations and anions produce diffusion potentials 220 How does membrane potential relate to ion distribution? 220 The Nernst equation distinguishes between active and'passive transport 221
Proton transport is a major determinant of the membrane potential 222 8.3 Membrane Transport Processes 223 Channels enhance diffusion across membranes 224 Carriers bind and transport specific substances 226 Primary active transport requires energy 226 Secondary active transport is driven by ion gradients 226 Kinetic analyses can elucidate transport mechanisms 228 8,4 Membrane Transport Proteins 228 190 7.1 Essential Nutrients, Deficiencies, and Plant Disorders 191 Special techniques are used in nutritional studies 193 Nutrient solutions can sustain rapid plant growth 194 Mineral deficiencies disrupt plant metabolism and function 195 Plant tissue analysis reveals mineral deficiencies 199 BOX 7.2 Root systems differ in form but are based on common structures 205 Different areas of the root absorb mineral ions differently 207 Nutrient availability influences root growth and development 208 Mycorrhizal symbioses facilitate nutrient uptake by roots 210 Nutrients move between mycorrhizal fungi and root cells 213 lonomics: A Powerful Approach to Study Mineral Nutrition 200 7.2 Treating Nutritional Deficiencies 201 Crop yields can be improved by the addition of fertilizers 201 Some mineral nutrients can be absorbed by leaves 202 7.3 Soil, Roots, and Microbes 202 Negatively charged soil particles affect the adsorption of mineral nutrients 202 Soil pH affects nutrient availability, soil microbes, and root growth 204 Excess m ineral ions in the soil limit plant growth 204 Some plants develop extensive root systems 205 Genes encoding many transporters have been identified 230
Transporters exist for diverse nitrogen containing compounds 230 Cation transporters are diverse 231 Anion transporters have been identified 233 Transporters for metal and metalloid ions transport essential micronutrients 234 Aquaporins have diverse functions 235 Plasma membrane H+-ATPases are highly regulated P-type ATPases 236 The tonoplast H+-ATPase drives solute accumulation in vacuoles 237 H+-pyrophosphatases and P-type H+-ATPases also pump protons at the tonoplast 238 8.5 Transport in Stomatai Guard Cells 238 Blue light induces stomatal opening 239 Abscisic acid and high CO2 induce stomatal closing 240 8.6 Ion Transport in Roots 240 Solutes move through both apoplast and symplasm 240 Ions cross both symplasm and apoplast 241 Xylem parenchyma cells participate in xylem loading 242
Table of Contents CHAPTER 9 ■ Photosynthesis: The Light Reactior 9.1 Photosynthesis in Green Plants 247 9,2 General Concepts 248 Light consists of photons with characteristic energies 248 Absorption of photosynthetically active light changes the electronic states of chlorophylls 248 Photosynthetic pigments absorb the light that powers photosynthesis 250 9.3 Key Experiments in Understanding Photosynthesis 252 Action spectra relate light absorption to photosynthetic activity 252 Photosynthesis takes place in complexes containing light-harvesting antennas and photochemical reaction centers 253 The chemical reaction of photosynthesis is driven by light 254 Light drives the reduction of NADW and the formation of ATP 254 Oxygen-evolving organisms have two photosystems that operate in series 255 9.4 Organization of the Photosynthetic Apparatus 256 The chloroplast is the site of photosynthesis 256 Thylakoids contain integral membrane proteins 257 Photosystems I and II are spatially separated in the thylakoid membrane 257 Anoxygenic photosynthetic bacteria have a single reaction center 259 9.5 Organization of Light-Absorbing Antenna Systems 259 Antenna systems contain chlorophyll and are membrane-associated 259 The antenna funnels energy to the reaction center 260 Many antenna pigment-protein complexes have a common structural motif 260 9.6 Mechanisms of Electron Transport 261 Electrons from chlorophyll travel through the carriers organized in the Z scheme 261 Energy is captured when an excited chlorophyll reduces an electron acceptor molecule 263 The reaction center chlorophylls of
the two photosystems absorb at different wavelengths 264 The PSII reaction center is a multi-subunit pigmentprotein complex 264 Water is oxidized to oxygen by PSII 264 Pheophytin and two quinones accept electrons from PSII 265 Electron flow through the cytochrome b6/complex also transports protons 266 xix Plastocyanin carries electrons between the cytochrome b¿f complex and photosystem I 268 The PSI reaction center oxidizes PC and reduces ferredoxin, which transfers electrons to NADW 268 Some herbicides block photosynthetic electron flow 269 9.7 Proton Transport and ATP Synthesis in the Chloroplast 270 Cyclic electron flow augments the output of ATP to balance the chloroplast energy budget 272 9.8 Repair and Regulation of the Photosynthetic Machinery 273 Carotenoids serve as photoprotective agents 273 Some xanthophylls also participate in energy dissipation 274 The PSII reaction center is easily damaged | | I and rapidly repaired 274 Thylakoid stacking permits energy partitioning between the photosystems 275 9.9 Genetics, Assembly, and Evolution of Photosynthetic Systems 275 Chloroplast genes exhibit non-Mendelian patterns of inheritance 275 Most chloroplast proteins are imported from the cytoplasm 275 ' The biosynthesis and breakdown of chlorophyll are complex pathways 276 Complex photosynthetic organisms have evolved from, simpler forms 276 CHAPTER 10 ’ f I I I | | ' lotosynthesis: The Carbon w . Reactions 281 I 10.1 The Calvin-Benson Cycle ■ 282 The Calvin-Benson cycle has three phases: carboxylation, reduction, and regeneration 282 The fixation of CO2 via carboxylation
of ribulose 1,5-bisphosphate and the reduction of 3-phosphoglycerate yield triose phosphates 283, The regeneration of ribulose 1,5-bisphosphate ensures the continuous assimilation of CO2 284 An induction period precedes the steady state of photosynthetic CO2 assimilation 285 Many mechanisms regulate the Calvin-Benson cycle 286 Rubisco activase regulates the catalytic activity of Rubisco 287 Light regulates the Calvin-Benson cycle via the ferredoxin-thioredoxin system 288 Light-dependent ion movements modulate enzymes of the Calvin-Benson cycle 289 Light controls the assembly of chloroplast enzymes into supramolecular complexes 289
xx Table of Contents 10.2 The Oxygenation Reaction of Rubisco and Photorespiration 290 The oxygenation of ribulose 1,5-bisphosphate sets in motion photorespiration 291 Photorespiration is linked to the photosynthetic electron transport system 295 Enzymes of plant photorespiration derive from different ancestors 295 BOX 10.1 Production of Biomass May Be Enhanced by Engineering Photorespiration 296 Photorespiration interacts with many metabolic pathways 296 10.3 Inorganic Carbon-Concentrating Mechanisms 297 10.4 Inorganic Carbon-Concentrating Mechanisms: C4 Photosynthetic Carbon Fixation 297 Malate and aspartate are the primary carboxylation products of the Cį cycle 298 Kranz-type C4 plants assimilate CO2 by the concerted action of two different types of cells 299 The C4 subtypes use different mechanisms to decarbox ylate four-carbon acids transported to bundle sheath cells 301 Bundle sheath cells and mesophyll cells exhibit anatomical and biochemical differences 301 The C4 cycle also concentrates CO2 in single cells 302 Light regulates the activity of key C4 enzymes 302 Photosynthetic assimilation of CO2 in C4 plants requires more transport processes than in C3 plants 302 In hot, dry climates, the C4 cycle reduces photorespiration 303 10.5 Inorganic Carbon-Concentrating Mechanisms: Crassulacean Acid Metabolism (CAM) 303 Different mechanisms regulate C4 PEPCase and CAMPEPCase 305 CAM is a versatile mechanism sensitive to environmental stimuli 305 10.6 Accumulation and Partitioning of Photosynthates—Starch and Sucrose 305 10.7 Formation and Mobilization of Chloroplast Starch
306 Chloroplast stroma accumulates starch as insoluble granules during the day 307 Starch degradation at night requires the phosphorylation of amylopectin 310 The export of maltose prevails in the nocturnal break down of transitory starch 310 The synthesis and degradation of the starch granule are regulated by multiple mechanisms 311 10.8 Sucrose Biosynthesis and Signaling 312 Triose phosphates from the Calvin-Benson cycle build up the cy tosolic pool of three important hexose phosphates in the light 312 Fructose 2,6-bisphosphate regulates the hexose phosphate pool in the light 314 Sucrose is continuously synthesized in the cy tosol 314 Sucrose plays only a minor role in stomatai regulation 316 CHAPTER 11 lotosynthesis: Physiological and Ecological Considerations 321 11.1 Photosynthesis Is Influenced by Leaf Properties 322 Leaf anatomy and canopy structure optimize light absorption 323 Leaf angle and leaf movement can control light absorption 325 Leaves acclimate to sun and shade environments 325 11.2 Effects of Light on Photosynthesis in the Intact Leaf 326 ■ Photosynthetic light-response curves reveal differences in leaf properties 326 Leaves must dissipate excess light energy as heat 328 Absorption of too much light can lead to photoinhibition 330 11.3 . Effects of Temperature on Photosynthesis in the Intact Leaf 331 Leaves mus t dissipate vast quantities of heat 331 There is an optimal temperature for photosynthesis 332 Photosynthesis is sensitive to both high and low temperatures 332 Photosynthetic efficiency is temperature-sensitive 333 11.4 Effects of Carbon Dioxide
on Photosynthesis in the Intact Leaf 334 Atmospheric CO2 concentration keeps rising 334 CO2 diffusion to the chloroplast is essential to photosynthesis 334 CO2 supply imposes limitations on photosy nthesis 336 How will photosynthesis and respiration change in the future under elevated CO2 conditions? 338 11.5 Stable Isotopes Record Photosynthetic Properties 340 How do we measure the stable carbon isotopes of plants? 340 Why does the carbon isotope ratio vary in plants? 341 CHAPTER 12 . anslocation in the Phloem 345 12.1 Patterns of Translocation: Source to Sink 346 12.2 Pathways of Translocation 347 Sugar is translocated in phloem sieve elements 347 Mature sieve elements are living cells specialized for translocation 347 Large pores in cell walls are the prominent feature of sieve elements 348 Companion cells aid the highly specialized sieve elements 350
Table of Contents 12.3 Phloem Loading 351 Phloem loading can occur via the apoplast or symplasm 351 Apoplastic loading is characteristic of many herbaceous species 352 Sucrose loading in the apoplastic pathway requires metabolic energy 352 Phloem loading in the apoplastic pathway involves a sucrose-H+ symporter 353 Transfer cells are companion cells that are specialized for membrane transport 353 Phloem loading is symplasmic in some species 354 The oligomer-trapping model explains symplasmic loading in plants with intermediary-type companion cells 354 Phloem loading is passive in several tree species 355 The type of phloem loading is correlated with several significant characteristics 356 12.4 Long-Distance Transport: A Pressure-Driven Mechanism 357 Mass transfer is much faster than diffusion 357 The pressure-flow model is a passive mechanism for phloem transport 357 The pressure is osmotically generated 357 Some predictions of pressure: flow have been confirmed, while others require further experimentation 359 Functional sieve plate pores appear to be open channels 359 Are the pressure gradients in the sieve elements sufficient to drive phloem transport in trees? 360 Modified models for translocation by mass flow have been suggested 361 Does translocation in gymnosperms involve a different mechanism? 361 12.5 Materials Translocated in the Phloem 361 Sugars are translocated in a nonreducing form 362 , Other small organic solutes are translocated in the phloem 362 Phloem-mobile macromolecules often originate in companion cells 364 Damaged sieve elements are sealed off 364
12.6 Phloem Unloading and Sink-to-Source Transition 365 Phloem unloading and short-distance transport can occur via symplasmic or apoplastic pathways 366 Symplasmic unloading supplies growing vegetative sinks 366 Symplasmic unloading is passive but depends on energy consumption in the sink 367 Import into seeds, fruits, and storage organs often involves an apoplastic step 367 Apoplastic import is active and requires metabolic energy 368 The transition of a leaf from sink to source is gradual 369 12.7 Photosynthate Distribution: Allocation and Partitioning 371 Allocation includes storage, utilization, and transport 371 xxi Source leaves regulate allocation 371 Various sinks partition transport sugars 372 Sink tissues compete for available translocated photosynthate 372 Sink strength depends on sink size and activity 372 The source adjusts over the long term to changes in the source-to-sink ratio 373 12.8 Transport of Signaling Molecules 373 Turgor pressure and chemical signals coordinate source and sink activities 374 Mobile RNAs function as signal molecules in the phloem to regulate growth and development 374 Mobile proteins also function as signal molecules to regulate growth and development 375 Plasmodesmata function in phloem, signaling 375 BOX 12.1 Relevance of Phloem Translocation and Signaling for Climate Change and Biotechnology 376 CHAPTER 13 ՛ I I •spiration and Lipid a Metabolism 379 । m I I I . i । l Հ 13.1 Overview of Plant Respiration 379 13.2 Glycolysis 382 I । Glycolysis metabolizes carbohydrates from several sources 382 The energy-conserving phase of
glycolysis produces pyruvate, ATP, and NADH 384 Plants have alternative glycolytic reactions 385 In the absence of oxygen, fermentation regenerates the NAD+ needed for glycolytic ATP production 385 13.3 The Oxidative Pentose Phosphate Pathway 386 The oxidative pentose phosphate pathway produces NADPH and biosynthetic intermediates 386 The oxidative pentose phosphate pathway is controlled by cellular redox status 388 13.4 The Tricarboxylic Acid Cycle 388 Mitochondria are semiautonomous organelles 388 Pyruvate enters the mitochondrion and is oxidized via the TCA cycle 389 The TCA cycle of plants has unique features 391 13.5 Oxidative Phosphorylation 391 The electron transport chain catalyzes a flow of electrons from NADH to O2 392 The electron transport chain has supplementary branches 393 ATP synthesis in the mitochondrion is coupled to electron transport 394 Transporters exchange substrates and products 396 Aerobic respiration yields about 60 molecules of ATP per molecule of sucrose 396 Several subunits of respiratory complexes are encoded by the mitochondrial genome 396 Plants have several mechanisms that lower the ATP yield 398
xxii Table of Contents Respiration is an integral part of a redox and biosynthesis network 400 Respiration is controlled at multiple levels 401 Free-living and symbiotic bacteria fix nitrogen 428 Nitrogen fixation requires microanaerobic or anaerobic conditions 429 BOX 14.1 13.6 Respiration in Intact Plants and Tissues 402 Plants respire roughly half of the daily photosynthetic yield 402 Respiratory processes operate during photosynthesis 403 Different tissues and organs respire at different rates 403 BOX 13.1 Symbiotic nitrogen fixation occurs in specialized structures 430 Establishing symbiosis requires an exchange of signals 431 Nod factors produced by bacteria act as signals for symbiosis 431 Nodule formation involves phytohormones 432 The nitrogenase enzyme complex fixes N2 434 Amides and ureides are the transported forms of nitrogen 435 Modifying Respiration for Future Needs 404 Environmental factors alter respiration rates 404 13.7 Lipid Metabolism 405 Fats and oils store large amounts of energy 405 Triacylglycerols are stored in oil bodies 406 BOX 13.2 Biotechnology of Lipids in a Changing World 407 Polar glycerolipids are the main structural lipids in membranes 407 Fatty acid biosynthesis consists of cycles of two-carbon addition 407 Glycerolipids are synthesized in the plastids and the ER 410 Lipid composition influences membrane function 411 Membrane lipids are precursors of important signaling compounds 411 Storage lipids are converted into carbohydrates in germinating seeds 411 14.6 Sulfur Assimilation 435 · Sulfate is the form of sulfur transported into plants
435 Sulfate assimilation requires the reduction of sulfate to cysteine 436 Sulfate assimilation occurs mostly in leaves 438 Methionine is synthesized from cysteine 438 14.7 Phosphate Assimilation 438 miRNAs contribute to phosphate and sulfate signaling 438 14.8 Oxygen Assimilation 439 14.9 The Energetics of Nutrient Assimilation 439 CHAPTER 15 CHAPTER 14 * ssimilation of Inorganic Nutrients 417 14.1 Nitrogen in the Environment Challenges and Solutions for Solving Nitrogen Deficiency in Future Agriculture 430 418 Nitrogen passes through several forms in a biogeo chemical cycle 418 Unassimilated ammonium or nitrate I may be dangerous 419 14.2 Nitrate Assimilation 420 Many factors regulate nitrate reductase 421 Nitrite reductase converts nitrite to ammonium 421 Both roots and shoots assimilate nitrate 422 Nitrate can be transported in both xylem and phloem 422 Transceptor contributes to nitrate signaling 423 14.3 Ammonium Assimilation 424 Converting ammonium to amino acids requires two enzymes 424 Ammonium can be assimilated via an alternative pathway 426 Transamination reactions transfer nitrogen 426 Asparagine and glutamine link carbon and nitrogen metabolism 426 14.4 Amino Acid Biosynthesis 426 14.5 Biological Nitrogen Fixation 427 )iot։c Stress -՛·· ’ 15.1 Defining Plant Stress 444 Physiological adjustment to abiotic stress involves trade-offs between vegetative and reproductive development 445 15.2 Acclimation and Adaptation 445 Adaptation to stress involves genetic modification over many generations 445 Acclimation allows plants to respond to environmental fluctuations
446 15.3 Environmental Factors and Their Biological Impacts on Plants 446 Water deficit decreases turgor pressure, increases ion toxicity, and inhibits photosynthesis 447 Temperature stress affects a broad spectrum of physiological processes 447 Flooding results in anaerobic stress to the root 448 Salinity stress has both osmotic and cytotoxic effects 449 During freezing stress, extracellular ice crystal formation causes cell dehydration 449 Heavy metals can both mimic essential mineral nutrients and generate ROS 449 Ozone and ultraviolet light generate ROS that cause lesions and induce PCD 450 Combinations of abiotic stresses can induce unique signaling and metabolic pathways 450
Table of Contents xxiii Interactions occur between abiotic and biotic stresses 451 Sequential exposure to different abiotic stresses some times confers cross-protection 451 Beneficial microbes can improve plant tolerance to abiotic stress 451 15.4 Stress-Sensing Mechanisms in Plants 452 Early-acting stress sensors provide the initial signal for the stress response 452 15.5 Signaling Pathways Activated in Response to Abiotic Stress 453 The signaling intermediates of many stress-response pathways can interact 453 Acclimation to stress involves transcriptional regulatory networks called régulons 455 Chloroplasts and mitochondria respond to abiotic stress by sending stress signals to the nucleus 456 Plant-wide waves of Ca2+ and ROS mediate systemic acquired acclimation 456 Epigenetic mechanisms, retrotransposons, and small RNAs provide additional protection against stress 456 Hormonal interactions regulate abiotic stress responses 459 CHAPTER 16' ■ Signals from Sunlight 475 16.1 Plant Photoreceptors 476 ՛ Photoresponses are driven by light quality or spectral properties of the energy absorbed 477 Plants responses to light can be distinguished by the amount of light required 478 16.2 Phytochromes 480 Phytochrome is the primary photoreceptor for red and far-red light 480 Phytochrome can interconvert between Pr and Pfr forms 480 Pfr is the physiologically active form of phytochrome 481 The phytochrome chromophore and protein both undergo conformational changes in response to red light 481 Pfr is partitioned between the cytosol and the nucleus 483 16.3 Phytochrome Responses 484
Phytochrome responses vary in lag time and escape time 484 Phytochrome responses fall into three main categories based on the amount of light required 484 Phytochrome A mediates responses to continuous far-red light 486 15.6 Physiological and Developmental A Mechanisms That Protect Plants against Abiotic Stress 460 Plants adjust osmotically to drying soils by accumulating solutes 460 Submerged organs develop aerenchyma tissue in response to hypoxia 461 Antioxidants and ROS-scavenging pathways protect cells from oxidative stress 462 Molecular chaperones and molecular shields protect proteins and membranes during abiotic stress 462 Plants can alter their membrane lipids in response to temperature and other abiotic stresses 463 Exclusion and internal tolerance mechanisms allow plants to cope with toxic ions 464 Phytochelatins and other chelators contribute to internal tolerance of toxic metal ions 465 Plants use cryoprotectant molecules and antifreeze proteins to prevent ice crystal formation 466 ABA signaling during water stress causes the massive efflux of K+ and anions from guard cells 466 Plants can alter their morphology in response to abiotic stress 467 The process of recovery from stress can be dangerous to the plant and requires a coordinated adjustment of plant metabolism and physiology 469 Phytochrome В mediates responses to continuous red or white light 486 Roles for phytochromes C, D, and E are emerging 486 16.4 Phytochrome Signaling Pathways 487 Phytochrome regulates membrane potentials and ion fluxes 487 Phy tochrome regulates gene expression 487 Phytochrome
interacting factors (PIFs) act early in signaling 488 ' . Phytochrome signaling involves protein phosphorylation and déphosphorylation 488 Phytochrome-induced photomorphogenesis involves pro tein degradation 489 16.5 Blue-Light Responses and Photoreceptors 490 . A Blue-light responses have characteristic kinetics and lag times 490 16.6 Cryptochromes 491 The activated FAD chromophore of cryptochrome causes a conformational change in the protein 491 cryl and cry2 have different developmental effects 492 Nuclear cryptochromes inhibit COPl-induced protein degradation 493 Cryptochrome can also bind to transcriptional regulators directly 493
xxiv Table of Contents 16.7 Interactions of Cryptochrome with Other Photoreceptors 493 Stem elongation is inhibited by both red and blue photoreceptors 493 Phytochrome interacts with cryptochrome to regulate flowering 494 The circadian clock is regulated by multiple aspects of light 494 16.8 Phototropins 495 Blue light induces changes in FMN absorption maxima associated with conformation changes 495 The LOV2 domain is primarily responsible for kinase activation in response to blue light 496 Blue light induces a conformational change that "uncages" the kinase domain of phototropin and leads to autophosphorylation 496 Phototropins trigger plant movements that enhance light use 496 Blue light initiates stomatai opening via activation of the plasma membrane H+-ATPase 498 16.9 Responses to Ultraviolet Radiation 500 CHAPTER 17 ■ Seed Dormancy, Germination, and Seedling Establishment 505 17.1 Seed Structure 506 Seed anatomy varies widely among different plant groups 506 17.2 Seed Dormancy 508 There are two basic types of seed dormancy mechanisms: exogenous and endogenous 508 Non-dormant seeds can exhibit vivipary and precocious germination 509 The ABA:GA ratio is the primary determinant of embry onic seed dormancy 510 17.3 Release from Dormancy 511 Light is an important signal that breaks dormancy in small seeds 511 Some seeds require either chilling or after-ripening to break dormancy 511 Seed dormancy can be broken by various chemical compounds 512 17.4 Seed Germination 512 Germination and postgermination can be divided into three phases corresponding to the phases of water
uptake 513 17.5 Mobilization of Stored Reserves 514 Cereal seeds are a model for understanding starch mobilization 515 Legume seeds are a model for understanding protein mobilization 516 Oilseeds are a model for understanding lipid remobilization 517 17.6 Seedling Growth and Establishment 517 The development of emerging seedlings is strongly influenced by light 517 Gibberellins and brassinosteroids both suppress photomorphogenesis in darkness 518 Hook opening is regulated by phytochrome, auxin, and ethylene 519 Vascular differentiation begins during seedling emergence 520 The root tip has specialized cells 520 Ethylene and other hormones regulate root hair development 521 17.7 Differential Growth Enables Successful Seedling Establishment 522 Ethylene affects microtubule orientation and induces lateral cell expansion 523 Auxin promotes growth in stems and coleoptiles, while inhibiting growth in roots 524 The minimum lag time for auxin-induced elongation is 10 minutes 525 Auxin-induced proton extrusion loosens the cell wall 526 17.8 Tropisms: Growth in Response to Directional Stimuli 526 Gravitropism involves the lateral redistribution •af auxin 526 The gravitropic stimulus perturbs the symmetric move ments of auxin 526 Gravity perception is triggered by the sedimentation of amyloplasts 529 Gravity sensing may involve pH and calcium ions (Ca2+) as second messengers 532 Thigmotropism involves signaling by Ca2+, pH, and reac tive oxygen species 533 Hydrotropism involves ABA signaling and asymmetric cytokinin responses 534 Phototropins are the light receptors involved in
phototropism 535 Phototropism is mediated by the lateral redistribution of auxin 535 Shoot phototropism occurs in a series of steps 536 CHAPTER 18 ■ Vegetative Growth and Organogenesis: Primary Growth of the Plant Axis 541 18.1 Meristematic Tissues: Foundations for Indeterminate Growth 541 The root and shoot apical meristems use similar strategies to enable indeterminate growth 542 18.2 The Root Apical Meristem 542 The root tip has four developmental zones 542 The origin of different root tissues can be traced to specific initial cells 543 Auxin and cytokinin contribute to the maintenance and function of the RAM 543 18.3 The Shoot Apical Meristem 545 The shoot apical meristem has distinct zones and layers 545 A combination of positive and negative interactions deter mines apical meristem size 546
xxv Table of Contents KNOX class homeodomain transcription factors help maintain proliferation in the SAM through regulation of cytokinin and GA concentrations 547 Localized auxin accumulation promotes leaf initiation 547 Axillary meristems form in the axils of leaf primordia 548 18.4 Leaf Development 549 Growth determines leaf shape 551 18.5 The Establishment of Leaf Polarity 551 A signal from the SAM initiates adaxial-abaxial polarity 551 Antagonism between sets of transcription factors determines adaxial-abaxial leaf polarity 552 MYB transcription factors, HD-ZIP HI proteins, and KN0X1 repression promote adaxial identity 552 Abaxial identity is determined by auxin, KANADI, and YABBY 552 Blade outgrowth is auxin dependent and regulated by the YABBY and WOX genes 553 Leaf proximal-distal polarity also depends on specific gene expression 553 . In compound leaves, de-repression of the KNOX1 gene promotes leaflet formation 554 18.6 Differentiation of Epidermal Cell Types 555 Guard cell identity is determined by a specialized epidermal lineage 555 Two groups of bHLH transcription factors govern stomata! cell identity transitions 556 Cell-to-cell peptide signals regulate stornata! patterning 557 Intrinsic polarity in the stomatai lineage aids stornata! spacing 557 Environmental factors also regulate stomata! density 558 Stomata development in monocots involves some genes that are orthologous to those in Arabidopsis 558 18.7 Venation Patterns in Leaves 559 The primary leaf vein is initiated discontinuously from the preexisting vascular system 560 Auxin canalization initiates
development of the leaf trace 560 Basipetal auxin transport from the LI layer of the leaf primordium initiates development of the leaf trace procambium 561 The existing vasculature guides the growth of the leaf trace 562 Vascular development proceeds from procambium differentiation 562 Higher-order leaf veins differentiate in a predictable hierarchical order 562 Auxin regulates higher-order vein formation and patterning 563 CHAPTER 19 I I ■ Vegetative Growth and Organogenesis: Branching and Secondary Growth 567 19.1 Shoot Branching and Architecture 568 Auxin, cytokinins, and strigolactones regulate axillary bud outgrowth 569 Auxin from the shoot tip maintains | apical dominance 569 Strigolactones act locally to repress axillary bud growth 571 I Cytokinins antagonize the effects of strigolactones 571 Integration of environmental and hormonal branching signals is required for plant fitness 572 Axillary bud dormancy is affected by season, position, and age factors 572 19.2 Root Branching and Architecture 573 Lateral root primordia arise from the xylem pole pericycle cells 573 Lateral root formation can be divided into four distinct stages 574 Lateral root founder cells undergo asymmetric cell divi sions to initiate formation of lateral root primordia 576 Monocots and eudicots differ in their predominant root types 576 Transcription factors regulate the gravitropic setpoint angles of lateral roots and shoots 577 Plants can modify their root system architecture to optimize water and nutrient uptake 578 19,3 Secondary Growth 578 Two types of lateral meristems are involved in
secondary growth 578 The vascular cambium produces secondary xylem and phloem 579 Mobile transcription factors pre-pattern the vascular cambium 580 ' The gene networks that control secondary meristems share similarities and differences with those that control the apical meristems 582 Several phytohormones regulate vascular cambium activity and differentiation of secondary xylem and phloem 584 The cork cambium gives rise to the outer corky layer called the periderm 585 Bark has diverse protective and storage functions 586 Epicormic buds covered by bark can sprout after forest fires 586 CHAPTER 20 ' I I | ■ The Control of Flowering and Floral Development 591 Հ I ' : 20.1 Floral Evocation: Integrating ■ Environmental Cues 591 ' 20.2 The Shoot Apex and Phase Changes 592 Plants progress through three developmental phases 592 Juvenile tissues are produced first and are located at the base of the shoot 592 Phase changes can be influenced by nutrients, gibberel lins, and other signals 593 20.3 Circadian Rhythms: The Clock Within 594 Circadian rhythms exhibit characteristic features 595 Phase shifting adjusts circadian rhythms to different day-night cycles 597 Phytochromes and cryptochromes entrain the clock 597
xxvi Table of Contents 20.4 Photoperiodism: Monitoring Day Length 597 Plants can be classified according to their photoperiodic responses 597 The leaf is the site of perception of the photoperiodic signal 599 The length of the night is important for floral induction 599 Night breaks can cancel the effec t of the dark period 600 Photoperiodic timekeeping during the night depends on a circadian clock 601 The external coincidence model is based on oscillating light sensitivity 602 The coincidence of CONSTANS expression and light promotes flowering in LDPs 602 SDPs use a coincidence mechanism to inhibit flowering in long days 603 BOX 20.1 Refining Molecular Mechanisms of Photoperiodic Flowering Happening in Natural Environments 604 Phytochrome is the primary photoreceptor in photoperiodism 605 A blue-light photoreceptor regulates flowering in some LDPs 606 20.5 Long-Distance Signaling Involved in Flowering 606 Grafting studies provided the first evidence for a. trans missible floral stimulus 607 Florigen is translocated in the phloem 608 20.6 The Identification of Florigen 608 The Arabidopsis protein FLOWERING LOCUS T (FT) is florigen 608 20.7 Vernalization: Promoting Flowering with Cold 610 Vernalization results in competence to flower at the shoot apical meristem 610 Vernalization can involve epigenetic changes in gene expression 611 A range of vernalization pathways may have evolved. 612 20.8 Multiple Pathways Involved in Flowering 612 Gibberellins and ethylene can induce flowering 612 The transition to flowering involves multiple factors and pathways 613 20.9 Floral
Meristems and Floral Organ Development 613 The shoot apical meristem in Arabidopsis changes with development 613 The four different types of floral organs are initiated as separate whorls 614 ' Two major categories of genes regulate floral development 615 Floral meristem identity genes regulate meristem function . 615 Homeotic mutations led to the identification of floral organ identity genes 616 The ABC model partially explains the determination of . floral organ identity 617 Arabidopsis Class E genes are required for the activities of the A, B, and C genes 618 According to the Quartet Model, floral organ identity is regulated by tetrameric complexes of the ABCE proteins 619 Class D genes are required for ovule formation 620 Floral asymmetry in flowers is regulated by gene expression 620 CHAPTER 21 ’ ’ ■ Sexual Reproduction: From Gametes to Fruits 625 21.1 Development of the Male and Female Gametophyte Generations 625 21.2 Formation of Male Gametophytes ' in the Stamen 627 Pollen grain formation occurs in two successive stages 627 The multilayered pollen cell wall is surprisingly complex 629 21.3 Female Gametophyte Development in the Ovule 630 The Arabidopsis gynoecium is an important model system for studying ovule development 631 The vast majority of angiosperms exhibit Polygonum-type embryo sac development 632 Functional megaspores undergo a series of free nuclear mitotic divisions followed by cellularization 632 21.4 Pollination and Fertilization in Flowering Plants 633 The progamic phase includes everything from pollen landing and tube growth to the fusion of sperm
and egg 633 Adhesion and hydration of a pollen grain on a compati ble flower depend on recognition between pollen and stigma surfaces 634 Ca2+֊triggered polarization of the pollen grain precedes tube formation 635 Pollen tubes grow by tip growth 636 Receptor-like kinases are thought to regulate the ROPI GTPase switch, a master regulator of tip growth 638 Pollen tube tip growth in the pistil is guided by both physical and chemical cues 638 Style tissue may condition pollen tubes to grow toward the embryo sac 639 Synergid cells release chemoattractants that guide pollen tube growth to the micropyle 640 Double fertilization occurs in three distinct stages 641 21.5 Selfing versus Outcrossing 642 Hermaphroditic and monoecious species have evolved floral features to ensure outcrossing 642 Cytoplasmic male sterility (CMS) occurs in the wild and is of great utility in agriculture 642 Self-incompatibility (SI) is the primary mechanism that enforces outcrossing in angiosperms 643
Table of Contents xxvii Two distinct genetic mechanisms govern self-incompatibility 644 The Brassicaceae sporophytic SI system is mediated by S locus-encoded receptors and ligands 645 Cytotoxic S-RNases and F-box proteins determine gametophytic self-incompatibility (GSI) 645 21.6 Apomixis: Asexual Reproduction by Seed . 647 Apomixis is not an evolutionary dead end 647 21.7 Endosperm Development 647 Cellularization of coenocytic endosperm in Arabidopsis progresses from the micropylar to the chalazal region 648 Cellularization of the coenocytic endosperm of cereals progresses centripetally 649 Endosperm development and embryogenesis can occur autonomously 650 Many of the genes that control endosperm development are differentially expressed maternal or paternal genes 651 Cells of the starchy endosperm and aleurone layer follow divergent developmental pathways 652 21.8 Seed Coat Development 652 Seed coat development appears to be regulated by the endosperm 652 21.9 Seed Maturation and Desiccation Tolerance 653 Seed filling and desiccation tolerance phases overlap in most species 654 The acquisition of desiccation tolerance involves many metabolic pathways 654 During the acquisition of desiccation tolerance, the cells of the embryo acquire aiglassy state 655 LEA proteins and nonreducing sugars have been impli cated in seed desiccation tolerance 655 Abscisic acid plays a key role in seed maturation 655 Coat-imposed dormancy is correlated with long-term ' seed viability 655 21.10 Fruit Development and Ripening 656 The phytohormones auxin and gibberellic acid (GA) regulate fruit
set and parthenocarpy 656 Specific transcription factors regulate the development of the dehiscence zone 658 Tomato is an important model system for studying fleshy fruit development 659 Fleshy fruits undergo ripening 660 Ripening involves changes in the color of fruit 660 Fruit softening involves the coordinated action of many cell wall-degrading enzymes 661 Taste and flavor reflect changes in acids, sugars, aroma, and other compounds 661 The causal link between ethylene and ripening was demonstrated in transgenic and mutant tomatoes 662 Climacteric and non-climacteric fruit differ in their ethylene responses 662 The ripening process is transcriptionally regulated 663 Studying the molecular mechanism of ripening can have commercial applications 664 CHAPTER 22 I I I I ■ Embryogenesis: The Origin of Plant Architecture 669 22.1 Embryogenesis in Monocots I I and Eudicots 670 I I . Embryogenesis differs between monocots and eudicots, but also shares common features 670 22.2 Establishment of Apical-Basal Polarity 672 Apical-basal polarity is established early in embryogenesis 672 Zygote polarization can be studied using live imaging 673 22.3 Mechanisms Guiding Embryogenesis 676 Intercellular signaling processes play key roles in guiding position-dependent development 677 Cell-cell communication during early embryo develop ment may be regulated by plasmodesmata 677 Mutant analyses have identified genes for signaling pro cesses that are essential for embryo organization 678 22.4 Auxin Signaling During Embryogenesis 680 Spatial patterns of auxin accumulation regulate key
developmental events 680 The GNOM protein establishes a polar distribution of PIN auxin efflux proteins 681 MONOPTEROS encodes a transcription factor that is activated by auxin 681 22,5 Radial Patterning During Embryogenesis 682 I | Procambial precursors for the vascular stele lie at the center of the radial axis 683 The differentiation of cortical and endodermal cells involves the intercellular movement of a transcription factor 684 22,6 Formation of the Root and Shoot Apical Meristems 686 Root formation involves MONOPTEROS and other auxin-regulated transcription factors 686 Shoot formation requires HD-ZIP III, SHOOT MERISTEMLESS, and WUSCHEL genes 687 Plants can initiate embryogenesis in multiple types of cells 687 CHAPTER 23 I I I I | ■ Plant Senescence and Developmental Cell Death '691 23.1 Programmed Cell Death 692 I I Distinct types of PCD occur in plants 693 Developmental PCD and pathogen-triggered PCD involve distinct processes 693 The autophagy pathway captures and degrades cellular constituents within lytic compartments 693 Autophagy plays a dual role in the regulation of plant PCD 695 Autophagy is required for nutrient recycling during plant senescence 696
xxviii Table of Contents 23.2 The Leaf Senescence Syndrome 696 Leaf senescence may be sequential, seasonal, or stress-induced 697 Leaves undergo massive structural and biochemical changes during leaf senescence 698 The autolysis of chloroplast proteins occurs in multiple compartments 698 The STAY-GREEN (SGR) protein is required for both LHCP II protein recycling and chlorophyll catabolism 699 23.3 Regulation of Leaf Senescence: A Multi-Layered Network 700 Leaf senescence depends on the comprehensive regulation of pathways that respond to endogenous and environ mental factors 701 Plant hormones and other signaling agents can act as positive or negative regulators of leaf senescence 706 Positive senescence regulators 707 Negative senescence regulators 708 23.4 Abscission 709 Organ abscission is regulated by developmental and environmental cues 711 23.5 Whole-Plant Senescence 713 Angiosperm life cycles may be annual, biennial, or perennial 714 Whole-plant senescence differs from aging in animals 714 The determinacy of shoot apical meristems is develop mentally regulated 715 Nutrient redistribution may trigger senescence in monocarpic plants 716 The productivity of tall trees continues to increase right up to the onset of senescence 716 CHAPTER 24 ■ Biotic Interactions 721 24.1 Plant Interactions with Beneficial Microorganisms 723 Nod factors are recognized by the Nod factor receptor (NFR) in legumes 723 Arbuscular mycorrhizal associations and nitrogen-fixing symbioses involve related signaling pathways 723 Rhizobacteria can increase nutrient availability, stimulate root
branching, and protect against pathogens 725 24.2 Herbivore Interactions That Harm Plants 725 Mechanical barriers provide a first line of defense against insect pests and pathogens 726 Plant specialized metabolites can deter insect herbivores 728 Plants store constitutive toxic compounds in specialized structures 728 Plants often store defense chemicals as nontoxic watersoluble sugar conjugates in the vacuole 730 24.3 Inducible Defense Responses to Insect . Herbivores 732 Plants can recognize specific components of insect saliva 733 Ca2+ signaling and activation of the MAP kinase pathway are early events associated with insect herbivory 734 Jasmonate activates defense responses against insect herbivores 734 Jasmonate acts through a conserved ubiquitin ligase signaling mechanism 735 Hormonal interactions contribute to planfoinsect herbi vore interactions 735 JA initiates the production of defense proteins that inhibit herbivore digestion 736 Herbivore damage induces systemic defenses 736 Glutamate receptor-like (GLR) genes are required for long-distance electrical signaling during herbivory 737 Herbivore-induced volatiles can repel herbivores and attract natural enemies 738 Herbivore-induced volatiles can serve as long-distance signals between plants 739 Herbivore-induced volatiles can also act as systemic ., signals within a plant 739 Defense responses to herbivores and pathogens are regulated by circadian rhythms 739 Insects have evolved mechanisms to defeat plant defenses 741 24.4 Plant Defenses against Pathogens 741 Microbial pathogens have evolved various strategies to
invade host plants 741 Pathogens produce effector molecules that aid in the colonization of their plant host cells 742 Plants can detect pathogens through perception of pathogen-derived "danger signals" 743 R genes provide resistance to individual pathogens by recognizing strain-specific effectors 744 The hypersensitive response is a common defense against pathogens 745 A single encounter with a pathogen may increase resistance to future attacks 746 The main components of the salicylic acid signaling pathway have been identified 746 Phytoalexins with antimicrobial activity accumulate after pathogen attack 747 RNA interference plays a central role in antiviral immune responses in plants 747 Some plant parasitic nematodes form specific associations through the formation of distinct feeding structures 748 Plants compete with other plants by secreting allelopathic specialized metabolites into the soil 749 Some plants are parasites of other plants 749 Glossary G-1 Illustration Credits IC-1 Index 1-1 |
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author2 | Taiz, Lincoln 1942- Møller, Ian M. 1950- Murphy, Angus Zeiger, Eduardo 1939- |
author2_role | edt edt edt edt |
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author_GND | (DE-588)141478993 (DE-588)141109068 (DE-588)1153851261 (DE-588)141479027 |
author_facet | Taiz, Lincoln 1942- Møller, Ian M. 1950- Murphy, Angus Zeiger, Eduardo 1939- |
building | Verbundindex |
bvnumber | BV048958941 |
classification_rvk | WN 1000 |
ctrlnum | (OCoLC)1385301948 (DE-599)BVBBV048958941 |
discipline | Biologie |
discipline_str_mv | Biologie |
edition | International seventh edition |
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genre_facet | Lehrbuch |
id | DE-604.BV048958941 |
illustrated | Illustrated |
index_date | 2024-07-03T21:59:54Z |
indexdate | 2024-10-04T10:00:35Z |
institution | BVB |
isbn | 9780197614204 |
language | English |
oai_aleph_id | oai:aleph.bib-bvb.de:BVB01-034222733 |
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physical | xxviii, 752, G-25, IC-8, I-50 Seiten Illustrationen, Diagramme |
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publisher | Oxford University Press |
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spelling | Plant physiology Plant physiology and development Lincoln Taiz (Professor emeritus, University of California, Santa Cruz, USA), Ian Max Møller (Associate Professor, Aarhus University, Denmark), Angus Murphy (Professor, University of Maryland), Eduardo Zeiger (Professor emeritus, University of California, Los Angeles, USA) International seventh edition New York ; Oxford Oxford University Press [2023] xxviii, 752, G-25, IC-8, I-50 Seiten Illustrationen, Diagramme txt rdacontent n rdamedia nc rdacarrier Entwicklungsphysiologie (DE-588)4152449-4 gnd rswk-swf Pflanzen (DE-588)4045539-7 gnd rswk-swf Pflanzenphysiologie (DE-588)4045580-4 gnd rswk-swf Plants / Development (DE-588)4123623-3 Lehrbuch gnd-content Pflanzenphysiologie (DE-588)4045580-4 s DE-604 Pflanzen (DE-588)4045539-7 s Entwicklungsphysiologie (DE-588)4152449-4 s Taiz, Lincoln 1942- (DE-588)141478993 edt Møller, Ian M. 1950- (DE-588)141109068 edt Murphy, Angus (DE-588)1153851261 edt Zeiger, Eduardo 1939- (DE-588)141479027 edt Digitalisierung UB Regensburg - ADAM Catalogue Enrichment application/pdf http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=034222733&sequence=000001&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA Inhaltsverzeichnis |
spellingShingle | Plant physiology and development Entwicklungsphysiologie (DE-588)4152449-4 gnd Pflanzen (DE-588)4045539-7 gnd Pflanzenphysiologie (DE-588)4045580-4 gnd |
subject_GND | (DE-588)4152449-4 (DE-588)4045539-7 (DE-588)4045580-4 (DE-588)4123623-3 |
title | Plant physiology and development |
title_alt | Plant physiology |
title_auth | Plant physiology and development |
title_exact_search | Plant physiology and development |
title_exact_search_txtP | Plant physiology and development |
title_full | Plant physiology and development Lincoln Taiz (Professor emeritus, University of California, Santa Cruz, USA), Ian Max Møller (Associate Professor, Aarhus University, Denmark), Angus Murphy (Professor, University of Maryland), Eduardo Zeiger (Professor emeritus, University of California, Los Angeles, USA) |
title_fullStr | Plant physiology and development Lincoln Taiz (Professor emeritus, University of California, Santa Cruz, USA), Ian Max Møller (Associate Professor, Aarhus University, Denmark), Angus Murphy (Professor, University of Maryland), Eduardo Zeiger (Professor emeritus, University of California, Los Angeles, USA) |
title_full_unstemmed | Plant physiology and development Lincoln Taiz (Professor emeritus, University of California, Santa Cruz, USA), Ian Max Møller (Associate Professor, Aarhus University, Denmark), Angus Murphy (Professor, University of Maryland), Eduardo Zeiger (Professor emeritus, University of California, Los Angeles, USA) |
title_short | Plant physiology and development |
title_sort | plant physiology and development |
topic | Entwicklungsphysiologie (DE-588)4152449-4 gnd Pflanzen (DE-588)4045539-7 gnd Pflanzenphysiologie (DE-588)4045580-4 gnd |
topic_facet | Entwicklungsphysiologie Pflanzen Pflanzenphysiologie Lehrbuch |
url | http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=034222733&sequence=000001&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA |
work_keys_str_mv | UT plantphysiology AT taizlincoln plantphysiologyanddevelopment AT møllerianm plantphysiologyanddevelopment AT murphyangus plantphysiologyanddevelopment AT zeigereduardo plantphysiologyanddevelopment |