Plant Physiology, Fifth Edition - Sinauer
Fifthifth EditiEditionditiLincoln TaizProfessor EmeritusUniversity of California, Santa CruzEduardo ZeigerProfessor EmeritusUniversity of California, Los AngelesSinauer Associates Inc., PublishersSunderland, Massachusetts U.S.A. Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd III 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:08:58 PM
ContentsCHAPTER 1Plant Cells 1Plant Life: Unifying Principles 2Overview of Plant Structure 2Plant cells are surrounded by rigid cell walls 2New cells are produced by dividingtissues called meristems 2Three major tissue systems make up theplant body 4Plant Cell Organelles 4Biological membranes are phospholipidbilayers that contain proteins 4The Endomembrane System 8The nucleus contains the majority of thegenetic material 8Gene expression involves both transcriptionand translation 10The endoplasmic reticulum is a networkof internal membranes 10Secretion of proteins from cells begins with therough ER (RER) 13Glycoproteins and polysaccharides destinedfor secretion are processed in the Golgiapparatus 14The plasma membrane has specialized regionsinvolved in membrane recycling 16Vacuoles have diverse functions in plant cells 16Independently Dividing Organelles Derivedfrom the Endomembrane System 17Oil bodies are lipid-storing organelles 17Microbodies play specialized metabolic roles inleaves and seeds 17Independently Dividing, SemiautonomousOrganelles 18Proplastids mature into specialized plastids indifferent plant tissues 21Chloroplast and mitochondrial division areindependent of nuclear division 21The Plant Cytoskeleton 22The plant cytoskeleton consists of microtubulesand microfilaments 22Microtubules and microfilaments can assembleand disassemble 23Cortical microtubules can move around the cell by“treadmilling” 24Cytoskeletal motor proteins mediate cytoplasmicstreaming and organelle traffic 24Cell Cycle Regulation 25Each phase of the cell cycle has a specific set ofbiochemical and cellular activities 26The cell cycle is regulated by cyclins andcyclin-dependent kinases 26Mitosis and cytokinesis involve both microtubulesand the endomembrane system 27Plasmodesmata 29Primary and secondary plasmodesmata help tomaintain tissue developmental gradients 29SUMMARY 31 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XVI 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:15 PM
CHAPTER 2Genome Organization and Gene Expression 35Nuclear Genome Organization 35The nuclear genome is packaged intochromatin 36Centromeres, telomeres, and nucleolar organizerscontain repetitive sequences 36Transposons are mobile sequences withinthe genome 37Polyploids contain multiple copies of the entiregenome 38Phenotypic and physiological responses topolyploidy are unpredictable 41Plant Cytoplasmic Genomes: Mitochondriaand Chloroplasts 42The endosymbiotic theory describes the originof cytoplasmic genomes 42Organellar genomes consist mostly of linearchromosomes 43Organellar genetics do not obeyMendelian laws 44Transcriptional Regulation of NuclearGene Expression 45RNA polymerase II binds to the promoterregion of most protein-coding genes 45UNIT IEpigenetic modifications help determine geneactivity 48Posttranscriptional Regulation ofNuclear Gene Expression 50RNA stability can be influenced bycis-elements 50Noncoding RNAs regulate mRNA activity viathe RNA interference (RNAi) pathway 50Posttranslational regulation determinesthe life span of proteins 54Tools for Studying Gene Function 55Mutant analysis can help to elucidategene function 55Molecular techniques can measure theactivity of genes 55Gene fusions can introduce reporter genes 56Genetic Modification of Crop Plants 59Transgenes can confer resistance toherbicides or plant pests 59Genetically modified organisms arecontroversial 60SUMMARY 61Transport and Translocation of Water and Solutes 65CHAPTER 3 Water and Plant Cells 67Water in Plant Life 67The Structure and Properties of Water 68Water is a polar molecule that forms hydrogenbonds 68Water is an excellent solvent 69Water has distinctive thermal properties relative toits size 69Water molecules are highly cohesive 69Water has a high tensile strength 70Diffusion and Osmosis 71Diffusion is the net movement of molecules byrandom thermal agitation 71Diffusion is most effective over short distances 72Osmosis describes the net movement of wateracross a selectively permeable barrier 73Water Potential 73The chemical potential of water represents thefree-energy status of water 74Three major factors contribute to cellwater potential 74Water potentials can be measured 75Water Potential of Plant Cells 75Water enters the cell along a water potentialgradient 75Water can also leave the cell in response to a waterpotential gradient 77Water potential and its components vary withgrowth conditions and location within theplant 77Cell Wall and Membrane Properties 78 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XVII 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:15 PM
XVIII TABLE OF CONTENTSSmall changes in plant cell volume cause largechanges in turgor pressure 78The rate at which cells gain or lose water isinfluenced by cell membrane hydraulicconductivity 79Aquaporins facilitate the movement of water acrosscell membranes 79CHAPTER 4Plant Water Status 80Physiological processes are affected by plant waterstatus 80Solute accumulation helps cells maintain turgorand volume 80SUMMARY 81Water Balance of Plants 85Water in the Soil 85A negative hydrostatic pressure in soil waterlowers soil water potential 86Water moves through the soil by bulk flow 87Xylem transport of water in trees faces physicalchallenges 94Plants minimize the consequences ofxylem cavitation 96Water Absorption by Roots 87Water moves in the root via the apoplast,symplast, and transmembrane pathways 88Solute accumulation in the xylem can generate“root pressure” 89Water Movement from the Leaf to theAtmosphere 96Leaves have a large hydraulic resistance 96The driving force for transpiration is thedifference in water vapor concentration 96Water loss is also regulated by the pathwayresistances 98Stomatal control couples leaf transpiration toleaf photosynthesis 98The cell walls of guard cells have specializedfeatures 99An increase in guard cell turgor pressureopens the stomata 101The transpiration ratio measures the relationshipbetween water loss and carbon gain 101Water Transport through the Xylem 90The xylem consists of two types of trachearyelements 90Water moves through the xylem bypressure-driven bulk flow 92Water movement through the xylem requiresa smaller pressure gradient than movementthrough living cells 93What pressure difference is needed to lift water100 meters to a treetop? 93The cohesion–tension theory explains water transport in the xylem 93Overview: The Soil–Plant–AtmosphereContinuum 102SUMMARY 102CHAPTER 5 Mineral Nutrition 107Essential Nutrients, Deficiencies,and Plant Disorders 108Special techniques are used in nutritionalstudies 110Nutrient solutions can sustain rapidplant growth 110Mineral deficiencies disrupt plant metabolismand function 113Analysis of plant tissues reveals mineraldeficiencies 117Treating Nutritional Deficiencies 117Crop yields can be improved by addition offertilizers 118Some mineral nutrients can be absorbed byleaves 118Soil, Roots, and Microbes 119Negatively charged soil particles affect the adsorption of mineral nutrients 119Soil pH affects nutrient availability, soil microbes,and root growth 120Excess mineral ions in the soil limit plantgrowth 120Plants develop extensive root systems 121Root systems differ in form but are based oncommon structures 121 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XVIII 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:15 PM
TABLE OF CONTENTSDifferent areas of the root absorb differentmineral ions 123Nutrient availability influences root growth 124Mycorrhizal fungi facilitate nutrient uptakeby roots 125XIXNutrients move from mycorrhizal fungi toroot cells 126SUMMARY 126CHAPTER 6 Solute Transport 131Passive and Active Transport 132Transport of Ions across MembraneBarriers 133Different diffusion rates for cations and anionsproduce diffusion potentials 134How does membrane potential relate to iondistribution? 134The Nernst equation distinguishes betweenactive and passive transport 136Proton transport is a major determinant ofthe membrane potential 137Membrane Transport Processes 137Channels enhance diffusion acrossmembranes 139Carriers bind and transport specific substances 140Primary active transport requires energy 140Secondary active transport uses storedenergy 142Kinetic analyses can elucidate transportmechanisms 143Membrane Transport Proteins 144UNIT IIThe genes for many transporters havebeen identified 144Transporters exist for diversenitrogen-containing compounds 146Cation transporters are diverse 147Anion transporters have been identified 148Metal transporters transport essentialmicronutrients 149Aquaporins have diverse functions 149Plasma membrane H -ATPases are highlyregulated P-type ATPases 150The tonoplast H -ATPase drives soluteaccumulation in vacuoles 151H -pyrophosphatases also pump protons atthe tonoplast 153Ion Transport in Roots 153Solutes move through both apoplast andsymplast 153Ions cross both symplast and apoplast 153Xylem parenchyma cells participate in xylemloading 154SUMMARY 156Biochemistry and Metabolism 161CHAPTER 7 Photosynthesis: The Light Reactions 163Photosynthesis in Higher Plants 164General Concepts 164Light has characteristics of both a particleand a wave 164When molecules absorb or emit light,they change their electronic state 165Photosynthetic pigments absorb the light thatpowers photosynthesis 166Key Experiments in UnderstandingPhotosynthesis 167Action spectra relate light absorption tophotosynthetic activity 168Photosynthesis takes place in complexescontaining light-harvesting antennas andphotochemical reaction centers 169The chemical reaction of photosynthesis isdriven by light 170Light drives the reduction of NADP and theformation of ATP 171Oxygen-evolving organisms have twophotosystems that operate in series 171Organization of the PhotosyntheticApparatus 172The chloroplast is the site of photosynthesis 172 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XIX 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:15 PM
XX TABLE OF CONTENTSThylakoids contain integral membrane proteins 173Photosystems I and II are spatially separatedin the thylakoid membrane 174Anoxygenic photosynthetic bacteria have asingle reaction center 174Organization of Light-AbsorbingAntenna Systems 176Antenna systems contain chlorophyll andare membrane associated 176The antenna funnels energy to thereaction center 176Many antenna pigment–protein complexeshave a common structural motif 176Mechanisms of Electron Transport 178Electrons from chlorophyll travel throughthe carriers organized in the “Z scheme” 178Energy is captured when an excited chlorophyllreduces an electron acceptor molecule 179The reaction center chlorophylls of the twophotosystems absorb at differentwavelengths 180The photosystem II reaction center is amultisubunit pigment–protein complex 181Water is oxidized to oxygen byphotosystem II 181Pheophytin and two quinones accept electronsfrom photosystem II 183Electron flow through the cytochrome b6fcomplex also transports protons 183Plastoquinone and plastocyanin carry electronsbetween photosystems II and I 184The photosystem I reaction centerreduces NADP 185Cyclic electron flow generates ATP but noNADPH 185Some herbicides block photosyntheticelectron flow 186Proton Transport and ATP Synthesisin the Chloroplast 187Repair and Regulation of thePhotosynthetic Machinery 189Carotenoids serve as photoprotective agents 190Some xanthophylls also participate in energydissipation 190The photosystem II reaction center is easilydamaged 191Photosystem I is protected from active oxygenspecies 191Thylakoid stacking permits energy partitioningbetween the photosystems 191Genetics, Assembly, and Evolution ofPhotosynthetic Systems 192Chloroplast genes exhibit non-Mendelian patternsof inheritance 192Most chloroplast proteins are imported fromthe cytoplasm 192The biosynthesis and breakdown of chlorophyllare complex pathways 192Complex photosynthetic organisms have evolvedfrom simpler forms 193SUMMARY 194CHAPTER 8 Photosynthesis: The Carbon Reactions 199The Calvin–Benson Cycle 200The Calvin–Benson cycle has three stages:carboxylation, reduction, and regeneration 200The carboxylation of ribulose 1,5-bisphosphate fixesCO2 for the synthesis of triose phosphates 201Ribulose 1,5-bisphosphate is regenerated forthe continuous assimilation of CO2 201An induction period precedes the steady stateof photosynthetic CO2 assimilation 204Regulation of the Calvin–Benson Cycle 205The activity of rubisco increases in the light 206Light regulates the Calvin–Benson cycle via theferredoxin–thioredoxin system 207Light-dependent ion movements modulate enzymes of the Calvin–Benson cycle 208Light controls the assembly of chloroplast enzymesinto supramolecular complexes 208The C2 Oxidative Photosynthetic CarbonCycle 208The carboxylation and the oxygenation of ribulose1,5-bisphosphate are competing reactions 210Photorespiration depends on the photosyntheticelectron transport system 213Photorespiration protects the photosynthetic apparatus under stress conditions 214Photorespiration may be engineered to increasethe production of biomass 214 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XX 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:15 PM
TABLE OF CONTENTSInorganic Carbon–ConcentratingMechanisms 216Inorganic Carbon–Concentrating Mechanisms:The C4 Carbon Cycle 216Malate and aspartate are carboxylation products ofthe C4 cycle 217Two different types of cells participate in the C4cycle 218The C4 cycle concentrates CO2 in the chloroplastsof bundle sheath cells 220The C4 cycle also concentrates CO2 in singlecells 221Light regulates the activity of key C4 enzymes 221In hot, dry climates, the C4 cycle reduces photorespiration and water loss 221Inorganic Carbon–Concentrating Mechanisms:Crassulacean Acid Metabolism (CAM) 221CAM is a versatile mechanism sensitive to environmental stimuli 223XXIFormation and Mobilization ofChloroplast Starch 225Starch is synthesized in the chloroplastduring the day 225Starch degradation at night requires thephosphorylation of amylopectin 228The export of maltose prevails in the nocturnalbreakdown of transitory starch 230Sucrose Biosynthesis and Signaling 231Triose phosphates supply the cytosolic poolof three important hexose phosphates in thelight 231Fructose 2,6-bisphosphate regulates the hexosephosphate pool in the light 235The cytosolic interconversion of hexose phosphates governs the allocation of assimilatedcarbon 235Sucrose is continuously synthesized in thecytosol 235SUMMARY 237Accumulation and Partitioning ofPhotosynthates—Starch and Sucrose 224CHAPTER 9Photosynthesis: Physiological and EcologicalConsiderations 243Photosynthesis Is the Primary Function ofLeaves 244Leaf anatomy maximizes light absorption 245Plants compete for sunlight 246Leaf angle and leaf movement can control lightabsorption 247Plants acclimate and adapt to sun and shadeenvironments 248Photosynthetic Responses to Light by theIntact Leaf 249Light-response curves reveal photosyntheticproperties 249Leaves must dissipate excess light energy 251Absorption of too much light can lead tophotoinhibition 253Photosynthetic Responses toTemperature 254Leaves must dissipate vast quantities of heat 254Photosynthesis is temperature sensitive 255There is an optimal temperature forphotosynthesis 256Photosynthetic Responses to CarbonDioxide 256Atmospheric CO2 concentration keeps rising 257CO2 diffusion to the chloroplast is essential tophotosynthesis 258Patterns of light absorption generate gradients ofCO2 fixation 259CO2 imposes limitations on photosynthesis 260How will photosynthesis and respiration change inthe future under elevated CO2 conditions? 261Identifying Different PhotosyntheticPathways 263How do we measure the stable carbon isotopes ofplants? 263Why are there carbon isotope ratio variations inplants? 264SUMMARY266 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XXI 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:15 PM
XXII TABLE OF CONTENTSCHAPTER 10 Translocation in the Phloem 271Pathways of Translocation 272Sugar is translocated in phloem sieveelements 273Mature sieve elements are living cells specializedfor translocation 273Large pores in cell walls are the prominent featureof sieve elements 274Damaged sieve elements are sealed off 274Companion cells aid the highly specialized sieveelements 276Patterns of Translocation: Source to Sink 276Materials Translocated in the Phloem 277Phloem sap can be collected and analyzed 278Sugars are translocated in nonreducing form 279Other solutes are translocated in the phloem 280Rates of Movement 280The Pressure-Flow Model, a PassiveMechanism for Phloem Transport 281An osmotically-generated pressure gradient drivestranslocation in the pressure-flow model 281The predictions of mass flow have beenconfirmed 282Sieve plate pores are open channels 283There is no bidirectional transport in single sieveelements 284The energy requirement for transport through thephloem pathway is small 284Positive pressure gradients exist in the phloemsieve elements 284Does translocation in gymnosperms involve adifferent mechanism? 285Phloem Loading 285Phloem loading can occur via the apoplast orsymplast 285Abundant data support the existence ofapoplastic loading in some species 286Sucrose uptake in the apoplastic pathwayrequires metabolic energy 286Phloem loading in the apoplastic pathwayinvolves a sucrose–H symporter 287Phloem loading is symplastic in somespecies 288The polymer-trapping model explainssymplastic loading in plants with intermediarycells 288Phloem loading is passive in a number oftree species 289The type of phloem loading is correlated with anumber of significant characteristics 290Phloem Unloading and Sink-to-SourceTransition 291Phloem unloading and short-distance transportcan occur via symplastic or apoplasticpathways 291Transport into sink tissues requires metabolicenergy 292The transition of a leaf from sink to source isgradual 292Photosynthate Distribution: Allocation andPartitioning 294Allocation includes storage, utilization, andtransport 294Various sinks partition transport sugars 295Source leaves regulate allocation 295Sink tissues compete for available translocatedphotosynthate 296Sink strength depends on sink size andactivity 296The source adjusts over the long term to changesin the source-to-sink ratio 297The Transport of Signaling Molecules 297Turgor pressure and chemical signals coordinatesource and sink activities 297Proteins and RNAs function as signal moleculesin the phloem to regulate growth anddevelopment 298SUMMARY299 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XXII 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:16 PM
TABLE OF CONTENTSXXIIICHAPTER 11 Respiration and Lipid Metabolism 305Overview of Plant Respiration 305Glycolysis 309Glycolysis metabolizes carbohydrates fromseveral sources 309The energy-conserving phase of glycolysisextracts usable energy 310Plants have alternative glycolytic reactions 310In the absence of oxygen, fermentationregenerates the NAD needed forglycolysis 311Plant glycolysis is controlled by its products 312The Oxidative Pentose PhosphatePathway 312The oxidative pentose phosphate pathwayproduces NADPH and biosyntheticintermediates 314The oxidative pentose phosphate pathway isredox-regulated 314The Citric Acid Cycle 315Mitochondria are semiautonomousorganelles 315Pyruvate enters the mitochondrion and isoxidized via the citric acid cycle 316The citric acid cycle of plants has uniquefeatures 317Mitochondrial Electron Transport andATP Synthesis 317The electron transport chain catalyzes a flow ofelectrons from NADH to O2 318The electron transport chain has supplementarybranches 320ATP synthesis in the mitochondrion is coupled toelectron transport 320Transporters exchange substrates andproducts 322Aerobic respiration yields about 60 moleculesof ATP per molecule of sucrose 322Several subunits of respiratory complexesare encoded by the mitochondrial genome 324Plants have several mechanisms that lowerthe ATP yield 324Short-term control of mitochondrialrespiration occurs at different levels 326Respiration is tightly coupled to otherpathways 327Respiration in Intact Plants and Tissues 327Plants respire roughly half of the dailyphotosynthetic yield 328Respiration operates during photosynthesis 329Different tissues and organs respire at differentrates 329Environmental factors alter respiration rates 329Lipid Metabolism 330Fats and oils store large amounts of energy 331Triacylglycerols are stored in oil bodies 331Polar glycerolipids are the main structural lipids inmembranes 332Fatty acid biosynthesis consists of cycles of twocarbon addition 334Glycerolipids are synthesized in the plastidsand the ER 335Lipid composition influences membranefunction 336Membrane lipids are precursors of importantsignaling compounds 336Storage lipids are converted into carbohydratesin germinating seeds 336SUMMARY 338 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XXIII 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:16 PM
XXIV TABLE OF CONTENTSCHAPTER 12 Assimilation of Mineral Nutrients 343Nitrogen in the Environment 344Nitrogen passes through several forms in a biogeochemical cycle 344Unassimilated ammonium or nitrate may be dangerous 346Nitrate Assimilation 346Many factors regulate nitrate reductase 347Nitrite reductase converts nitrite toammonium 347Both roots and shoots assimilate nitrate 348Ammonium Assimilation 348Converting ammonium to amino acids requirestwo enzymes 348Ammonium can be assimilated via analternative pathway 350Transamination reactions transfer nitrogen 350Asparagine and glutamine link carbon andnitrogen metabolism 350Amino Acid Biosynthesis 351Biological Nitrogen Fixation 351Free-living and symbiotic bacteria fixnitrogen 351Nitrogen fixation requires anaerobicconditions 352Symbiotic nitrogen fixation occurs inspecialized structures 354Establishing symbiosis requires an exchange ofsignals 354Nod factors produced by bacteria act as signalsfor symbiosis 354Nodule formation involves phytohormones 355The nitrogenase enzyme complex fixes N2 357Amides and ureides are the transportedforms of nitrogen 358Sulfur Assimilation 358Sulfate is the absorbed form of sulfur inplants 358Sulfate assimilation requires the reduction ofsulfate to cysteine 359Sulfate assimilation occurs mostly in leaves 360Methionine is synthesized from cysteine 360Phosphate Assimilation 360Cation Assimilation 361Cations form noncovalent bonds with carboncompounds 361Roots modify the rhizosphere to acquire iron 362Iron forms complexes with carbonand phosphate 363Oxygen Assimilation 363The Energetics of Nutrient Assimilation 364SUMMARY 365CHAPTER 13 Secondary Metabolites and Plant Defense 369Secondary Metabolites 370Secondary metabolites defend plants against herbivores and pathogens 370Secondary metabolites are divided into three major groups 370Terpenes 370Terpenes are formed by the fusion of five-carbonisoprene units 370There are two pathways for terpenebiosynthesis 370IPP and its isomer combine to form largerterpenes 371Some terpenes have roles in growth anddevelopment 373Terpenes defend many plants againstherbivores 373Phenolic Compounds 374Phenylalanine is an intermediate in thebiosynthesis of most plant phenolics 375Ultraviolet light activates some simplephenolics 377The release of phenolics into the soil maylimit the growth of other plants 377Lignin is a highly complex phenolicmacromolecule 377 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XXIV 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:16 PM
TABLE OF CONTENTSThere are four major groups of flavonoids 378Anthocyanins are colored flavonoids that attractanimals 378Flavones and flavonols may protect against damage by ultraviolet light 379Isoflavonoids have widespread pharmacologicalactivity 379Tannins deter feeding by herbivores 380Nitrogen-Containing Compounds 381Alkaloids have dramatic physiological effects onanimals 381Cyanogenic glycosides release the poisonhydrogen cyanide 384Glucosinolates release volatile toxins 385Nonprotein amino acids are toxic toherbivores 385Induced Plant Defenses against InsectHerbivores 386Plants can recognize specific components ofinsect saliva 386Jasmonic acid activates many defensiveresponses 387Some plant proteins inhibit herbivoredigestion 389UNIT IIIXXVDamage by insect herbivores inducessystemic defenses 389Herbivore-induced volatiles have complexecological functions 389Insects have developed strategies to copewith plant defenses 391Plant Defenses against Pathogens 391Pathogens have developed various strategies toinvade host plants 391Some antimicrobial compounds are synthesizedbefore pathogen attack 392Infection induces additional antipathogendefenses 392Phytoalexins often increase after pathogenattack 393Some plants recognize specific pathogen-derivedsubstances 393Exposure to elicitors induces a signal transductioncascade 394A single encounter with a pathogen may increaseresistance to future attacks 394Interactions of plants with nonpathogenic bacteriacan trigger induced systemic resistance 395SUMMARY 396Growth and Development 401CHAPTER 14 Signal Transduction 403Signal Transduction in Plant andAnimal Cells 404Plants and animals have similar transductioncomponents 404Receptor kinases can initiate a signaltransduction cascade 406Plants signal transduction components haveevolved from both prokaryotic and eukaryoticancestors 406Signals are perceived at many locationswithin plant cells 408Plant signal transduction often involvesinactivation of repressor proteins 409Protein degradation is a common feature inplant signaling pathways 411Several plant hormone receptors encodecomponents of the ubiquitination machinery 413Inactivation of repressor proteins results in agene expression response 414Plants have evolved mechanisms for switchingoff or attenuating signaling responses 414Cross-regulation allows signal transductionpathways to be integrated 416Signal Transduction in Space and Time 418Plant signal transduction occurs over a wide rangeof distances 418The timescale of plant signal transduction rangesfrom seconds to years 419SUMMARY 421 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.TAIZ FM JD.indd XXV 2012 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.5/19/10 4:09:16 PM
XXVI TABLE OF CONTENTSCHAPTER 15Cell Walls: Structure, Biogenesis,and Expansion 425The Structure and Synthesis ofPlant Cell Walls 426Plant cell walls have varied architecture 426The primary cell wall is composed ofcellulose microfibrils embedded in apolysaccharide matrix 428Cellulose microfibrils are synthesized atthe plasma membrane 430Matrix polymers are synthesized in theGolgi apparatus and secreted via vesicles 433Hemicelluloses are matrix polysaccharidesthat bind to cellulose 433Pectins are hydrophilic gel-forming componentsof the matrix 434Structural proteins become cross-linked int
Plant Life: Unifying Principles 2 Overview of Plant Structure 2 Plant cells are surrounded by rigid cell walls 2 . Mineral defi ciencies disrupt plant metabolism and function 113 Analysis of plant tissues reveals miner
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