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Fifth

ifth Editi
Edition
diti

Lincoln Taiz
Professor Emeritus
University of California, Santa Cruz

Eduardo Zeiger
Professor Emeritus
University of California, Los Angeles

Sinauer Associates Inc., Publishers


Sunderland, Massachusetts U.S.A.

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Contents

CHAPTER 1 Plant Cells 1


Plant Life: Unifying Principles 2 Independently Dividing, Semiautonomous
Organelles 18
Overview of Plant Structure 2
Proplastids mature into specialized plastids in
Plant cells are surrounded by rigid cell walls 2 different plant tissues 21
New cells are produced by dividing Chloroplast and mitochondrial division are
tissues called meristems 2 independent of nuclear division 21
Three major tissue systems make up the
plant body 4 The Plant Cytoskeleton 22
The plant cytoskeleton consists of microtubules
Plant Cell Organelles 4 and microfilaments 22
Biological membranes are phospholipid Microtubules and microfilaments can assemble
bilayers that contain proteins 4 and disassemble 23
The Endomembrane System 8 Cortical microtubules can move around the cell by
The nucleus contains the majority of the treadmilling 24
genetic material 8 Cytoskeletal motor proteins mediate cytoplasmic
Gene expression involves both transcription streaming and organelle traffic 24
and translation 10 Cell Cycle Regulation 25
The endoplasmic reticulum is a network Each phase of the cell cycle has a specific set of
of internal membranes 10 biochemical and cellular activities 26
Secretion of proteins from cells begins with the The cell cycle is regulated by cyclins and
rough ER (RER) 13 cyclin-dependent kinases 26
Glycoproteins and polysaccharides destined Mitosis and cytokinesis involve both microtubules
for secretion are processed in the Golgi and the endomembrane system 27
apparatus 14
The plasma membrane has specialized regions Plasmodesmata 29
involved in membrane recycling 16 Primary and secondary plasmodesmata help to
Vacuoles have diverse functions in plant cells 16 maintain tissue developmental gradients 29
Independently Dividing Organelles Derived SUMMARY 31
from the Endomembrane System 17
Oil bodies are lipid-storing organelles 17
Microbodies play specialized metabolic roles in
leaves and seeds 17

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CHAPTER 2 Genome Organization and Gene Expression 35
Nuclear Genome Organization 35 Epigenetic modifications help determine gene
The nuclear genome is packaged into activity 48
chromatin 36 Posttranscriptional Regulation of
Centromeres, telomeres, and nucleolar organizers Nuclear Gene Expression 50
contain repetitive sequences 36
RNA stability can be influenced by
Transposons are mobile sequences within cis-elements 50
the genome 37
Noncoding RNAs regulate mRNA activity via
Polyploids contain multiple copies of the entire the RNA interference (RNAi) pathway 50
genome 38
Posttranslational regulation determines
Phenotypic and physiological responses to the life span of proteins 54
polyploidy are unpredictable 41
Tools for Studying Gene Function 55
Plant Cytoplasmic Genomes: Mitochondria
Mutant analysis can help to elucidate
and Chloroplasts 42 gene function 55
The endosymbiotic theory describes the origin Molecular techniques can measure the
of cytoplasmic genomes 42 activity of genes 55
Organellar genomes consist mostly of linear Gene fusions can introduce reporter genes 56
chromosomes 43
Organellar genetics do not obey Genetic Modification of Crop Plants 59
Mendelian laws 44 Transgenes can confer resistance to
herbicides or plant pests 59
Transcriptional Regulation of Nuclear
Gene Expression 45 Genetically modified organisms are
controversial 60
RNA polymerase II binds to the promoter
region of most protein-coding genes 45 SUMMARY 61

UNIT I Transport and Translocation of Water and Solutes 65


CHAPTER 3 Water and Plant Cells 67
Water in Plant Life 67 Water Potential 73
The Structure and Properties of Water 68 The chemical potential of water represents the
free-energy status of water 74
Water is a polar molecule that forms hydrogen
bonds 68 Three major factors contribute to cell
water potential 74
Water is an excellent solvent 69
Water potentials can be measured 75
Water has distinctive thermal properties relative to
its size 69 Water Potential of Plant Cells 75
Water molecules are highly cohesive 69 Water enters the cell along a water potential
Water has a high tensile strength 70 gradient 75
Water can also leave the cell in response to a water
Diffusion and Osmosis 71 potential gradient 77
Diffusion is the net movement of molecules by Water potential and its components vary with
random thermal agitation 71 growth conditions and location within the
Diffusion is most effective over short distances 72 plant 77
Osmosis describes the net movement of water
Cell Wall and Membrane Properties 78
across a selectively permeable barrier 73
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XVIII TABLE OF CONTENTS

Small changes in plant cell volume cause large Plant Water Status 80
changes in turgor pressure 78 Physiological processes are affected by plant water
The rate at which cells gain or lose water is status 80
influenced by cell membrane hydraulic Solute accumulation helps cells maintain turgor
conductivity 79 and volume 80
Aquaporins facilitate the movement of water across
cell membranes 79 SUMMARY 81

CHAPTER 4 Water Balance of Plants 85


Water in the Soil 85 Xylem transport of water in trees faces physical
A negative hydrostatic pressure in soil water challenges 94
lowers soil water potential 86 Plants minimize the consequences of
Water moves through the soil by bulk flow 87 xylem cavitation 96

Water Absorption by Roots 87 Water Movement from the Leaf to the


Water moves in the root via the apoplast, Atmosphere 96
symplast, and transmembrane pathways 88 Leaves have a large hydraulic resistance 96
Solute accumulation in the xylem can generate The driving force for transpiration is the
root pressure 89 difference in water vapor concentration 96
Water loss is also regulated by the pathway
Water Transport through the Xylem 90 resistances 98
The xylem consists of two types of tracheary Stomatal control couples leaf transpiration to
elements 90 leaf photosynthesis 98
Water moves through the xylem by The cell walls of guard cells have specialized
pressure-driven bulk flow 92 features 99
Water movement through the xylem requires An increase in guard cell turgor pressure
a smaller pressure gradient than movement opens the stomata 101
through living cells 93
The transpiration ratio measures the relationship
What pressure difference is needed to lift water between water loss and carbon gain 101
100 meters to a treetop? 93
The cohesiontension theory explains water trans- Overview: The SoilPlantAtmosphere
port in the xylem 93 Continuum 102
SUMMARY 102

CHAPTER 5 Mineral Nutrition 107


Essential Nutrients, Deficiencies, Some mineral nutrients can be absorbed by
and Plant Disorders 108 leaves 118
Special techniques are used in nutritional Soil, Roots, and Microbes 119
studies 110
Negatively charged soil particles affect the adsorp-
Nutrient solutions can sustain rapid tion of mineral nutrients 119
plant growth 110
Soil pH affects nutrient availability, soil microbes,
Mineral deficiencies disrupt plant metabolism and root growth 120
and function 113
Excess mineral ions in the soil limit plant
Analysis of plant tissues reveals mineral growth 120
deficiencies 117
Plants develop extensive root systems 121
Treating Nutritional Deficiencies 117 Root systems differ in form but are based on
Crop yields can be improved by addition of common structures 121
fertilizers 118

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TABLE OF CONTENTS XIX

Different areas of the root absorb different Nutrients move from mycorrhizal fungi to
mineral ions 123 root cells 126
Nutrient availability influences root growth 124 SUMMARY 126
Mycorrhizal fungi facilitate nutrient uptake
by roots 125

CHAPTER 6 Solute Transport 131


Passive and Active Transport 132 The genes for many transporters have
been identified 144
Transport of Ions across Membrane
Transporters exist for diverse
Barriers 133
nitrogen-containing compounds 146
Different diffusion rates for cations and anions
Cation transporters are diverse 147
produce diffusion potentials 134
Anion transporters have been identified 148
How does membrane potential relate to ion
distribution? 134 Metal transporters transport essential
micronutrients 149
The Nernst equation distinguishes between
active and passive transport 136 Aquaporins have diverse functions 149
Proton transport is a major determinant of Plasma membrane H+-ATPases are highly
the membrane potential 137 regulated P-type ATPases 150
The tonoplast H+-ATPase drives solute
Membrane Transport Processes 137 accumulation in vacuoles 151
Channels enhance diffusion across H+-pyrophosphatases also pump protons at
membranes 139 the tonoplast 153
Carriers bind and transport specific substances 140
Ion Transport in Roots 153
Primary active transport requires energy 140
Solutes move through both apoplast and
Secondary active transport uses stored
symplast 153
energy 142
Ions cross both symplast and apoplast 153
Kinetic analyses can elucidate transport
mechanisms 143 Xylem parenchyma cells participate in xylem
loading 154
Membrane Transport Proteins 144
SUMMARY 156

UNIT II Biochemistry and Metabolism 161


CHAPTER 7 Photosynthesis: The Light Reactions 163
Photosynthesis in Higher Plants 164 Photosynthesis takes place in complexes
containing light-harvesting antennas and
General Concepts 164 photochemical reaction centers 169
Light has characteristics of both a particle The chemical reaction of photosynthesis is
and a wave 164 driven by light 170
When molecules absorb or emit light, Light drives the reduction of NADP and the
they change their electronic state 165 formation of ATP 171
Photosynthetic pigments absorb the light that Oxygen-evolving organisms have two
powers photosynthesis 166 photosystems that operate in series 171
Key Experiments in Understanding Organization of the Photosynthetic
Photosynthesis 167 Apparatus 172
Action spectra relate light absorption to The chloroplast is the site of photosynthesis 172
photosynthetic activity 168

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XX TABLE OF CONTENTS

Thylakoids contain integral membrane proteins 173 The photosystem I reaction center
Photosystems I and II are spatially separated reduces NADP+ 185
in the thylakoid membrane 174 Cyclic electron flow generates ATP but no
Anoxygenic photosynthetic bacteria have a NADPH 185
single reaction center 174 Some herbicides block photosynthetic
electron flow 186
Organization of Light-Absorbing
Antenna Systems 176 Proton Transport and ATP Synthesis
Antenna systems contain chlorophyll and in the Chloroplast 187
are membrane associated 176 Repair and Regulation of the
The antenna funnels energy to the Photosynthetic Machinery 189
reaction center 176
Carotenoids serve as photoprotective agents 190
Many antenna pigmentprotein complexes
Some xanthophylls also participate in energy
have a common structural motif 176
dissipation 190
Mechanisms of Electron Transport 178 The photosystem II reaction center is easily
Electrons from chlorophyll travel through damaged 191
the carriers organized in the Z scheme 178 Photosystem I is protected from active oxygen
Energy is captured when an excited chlorophyll species 191
reduces an electron acceptor molecule 179 Thylakoid stacking permits energy partitioning
The reaction center chlorophylls of the two between the photosystems 191
photosystems absorb at different
Genetics, Assembly, and Evolution of
wavelengths 180
Photosynthetic Systems 192
The photosystem II reaction center is a
multisubunit pigmentprotein complex 181 Chloroplast genes exhibit non-Mendelian patterns
of inheritance 192
Water is oxidized to oxygen by
photosystem II 181 Most chloroplast proteins are imported from
the cytoplasm 192
Pheophytin and two quinones accept electrons
from photosystem II 183 The biosynthesis and breakdown of chlorophyll
are complex pathways 192
Electron flow through the cytochrome b6f
complex also transports protons 183 Complex photosynthetic organisms have evolved
from simpler forms 193
Plastoquinone and plastocyanin carry electrons
between photosystems II and I 184 SUMMARY 194

CHAPTER 8 Photosynthesis: The Carbon Reactions 199


The CalvinBenson Cycle 200 Light-dependent ion movements modulate en-
The CalvinBenson cycle has three stages: zymes of the CalvinBenson cycle 208
carboxylation, reduction, and regeneration 200 Light controls the assembly of chloroplast enzymes
The carboxylation of ribulose 1,5-bisphosphate fixes into supramolecular complexes 208
CO2 for the synthesis of triose phosphates 201 The C2 Oxidative Photosynthetic Carbon
Ribulose 1,5-bisphosphate is regenerated for Cycle 208
the continuous assimilation of CO2 201
The carboxylation and the oxygenation of ribulose
An induction period precedes the steady state 1,5-bisphosphate are competing reactions 210
of photosynthetic CO2 assimilation 204
Photorespiration depends on the photosynthetic
Regulation of the CalvinBenson Cycle 205 electron transport system 213
The activity of rubisco increases in the light 206 Photorespiration protects the photosynthetic ap-
Light regulates the CalvinBenson cycle via the paratus under stress conditions 214
ferredoxinthioredoxin system 207 Photorespiration may be engineered to increase
the production of biomass 214

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TABLE OF CONTENTS XXI

Inorganic CarbonConcentrating Formation and Mobilization of


Mechanisms 216 Chloroplast Starch 225
Starch is synthesized in the chloroplast
Inorganic CarbonConcentrating Mechanisms: during the day 225
The C4 Carbon Cycle 216
Starch degradation at night requires the
Malate and aspartate are carboxylation products of phosphorylation of amylopectin 228
the C4 cycle 217
The export of maltose prevails in the nocturnal
Two different types of cells participate in the C4 breakdown of transitory starch 230
cycle 218
The C4 cycle concentrates CO2 in the chloroplasts Sucrose Biosynthesis and Signaling 231
of bundle sheath cells 220 Triose phosphates supply the cytosolic pool
The C4 cycle also concentrates CO2 in single of three important hexose phosphates in the
cells 221 light 231
Light regulates the activity of key C4 enzymes 221 Fructose 2,6-bisphosphate regulates the hexose
phosphate pool in the light 235
In hot, dry climates, the C4 cycle reduces photo-
respiration and water loss 221 The cytosolic interconversion of hexose phos-
phates governs the allocation of assimilated
Inorganic CarbonConcentrating Mechanisms: carbon 235
Crassulacean Acid Metabolism (CAM) 221 Sucrose is continuously synthesized in the
CAM is a versatile mechanism sensitive to environ- cytosol 235
mental stimuli 223
SUMMARY 237
Accumulation and Partitioning of
PhotosynthatesStarch and Sucrose 224

Photosynthesis: Physiological and Ecological


CHAPTER 9
Considerations 243
Photosynthesis Is the Primary Function of There is an optimal temperature for
Leaves 244 photosynthesis 256
Leaf anatomy maximizes light absorption 245 Photosynthetic Responses to Carbon
Plants compete for sunlight 246 Dioxide 256
Leaf angle and leaf movement can control light Atmospheric CO2 concentration keeps rising 257
absorption 247 CO2 diffusion to the chloroplast is essential to
Plants acclimate and adapt to sun and shade photosynthesis 258
environments 248 Patterns of light absorption generate gradients of
Photosynthetic Responses to Light by the CO2 fixation 259
Intact Leaf 249 CO2 imposes limitations on photosynthesis 260
Light-response curves reveal photosynthetic How will photosynthesis and respiration change in
properties 249 the future under elevated CO2 conditions? 261
Leaves must dissipate excess light energy 251 Identifying Different Photosynthetic
Absorption of too much light can lead to Pathways 263
photoinhibition 253 How do we measure the stable carbon isotopes of
Photosynthetic Responses to plants? 263
Temperature 254 Why are there carbon isotope ratio variations in
Leaves must dissipate vast quantities of heat 254 plants? 264
Photosynthesis is temperature sensitive 255 SUMMARY 266

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XXII TABLE OF CONTENTS

CHAPTER 10 Translocation in the Phloem 271


Pathways of Translocation 272 Phloem loading in the apoplastic pathway
Sugar is translocated in phloem sieve involves a sucroseH+ symporter 287
elements 273 Phloem loading is symplastic in some
Mature sieve elements are living cells specialized species 288
for translocation 273 The polymer-trapping model explains
Large pores in cell walls are the prominent feature symplastic loading in plants with intermediary
of sieve elements 274 cells 288
Damaged sieve elements are sealed off 274 Phloem loading is passive in a number of
tree species 289
Companion cells aid the highly specialized sieve
elements 276 The type of phloem loading is correlated with a
number of significant characteristics 290
Patterns of Translocation: Source to Sink 276
Phloem Unloading and Sink-to-Source
Materials Translocated in the Phloem 277 Transition 291
Phloem sap can be collected and analyzed 278 Phloem unloading and short-distance transport
Sugars are translocated in nonreducing form 279 can occur via symplastic or apoplastic
Other solutes are translocated in the phloem 280 pathways 291
Transport into sink tissues requires metabolic
Rates of Movement 280 energy 292
The Pressure-Flow Model, a Passive The transition of a leaf from sink to source is
Mechanism for Phloem Transport 281 gradual 292
An osmotically-generated pressure gradient drives Photosynthate Distribution: Allocation and
translocation in the pressure-flow model 281 Partitioning 294
The predictions of mass flow have been Allocation includes storage, utilization, and
confirmed 282 transport 294
Sieve plate pores are open channels 283 Various sinks partition transport sugars 295
There is no bidirectional transport in single sieve Source leaves regulate allocation 295
elements 284
Sink tissues compete for available translocated
The energy requirement for transport through the photosynthate 296
phloem pathway is small 284
Sink strength depends on sink size and
Positive pressure gradients exist in the phloem activity 296
sieve elements 284
The source adjusts over the long term to changes
Does translocation in gymnosperms involve a in the source-to-sink ratio 297
different mechanism? 285
The Transport of Signaling Molecules 297
Phloem Loading 285
Turgor pressure and chemical signals coordinate
Phloem loading can occur via the apoplast or source and sink activities 297
symplast 285
Proteins and RNAs function as signal molecules
Abundant data support the existence of in the phloem to regulate growth and
apoplastic loading in some species 286 development 298
Sucrose uptake in the apoplastic pathway
requires metabolic energy 286 SUMMARY 299

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TABLE OF CONTENTS XXIII

CHAPTER 11 Respiration and Lipid Metabolism 305


Overview of Plant Respiration 305 Aerobic respiration yields about 60 molecules
of ATP per molecule of sucrose 322
Glycolysis 309
Several subunits of respiratory complexes
Glycolysis metabolizes carbohydrates from are encoded by the mitochondrial genome 324
several sources 309
Plants have several mechanisms that lower
The energy-conserving phase of glycolysis the ATP yield 324
extracts usable energy 310
Short-term control of mitochondrial
Plants have alternative glycolytic reactions 310 respiration occurs at different levels 326
In the absence of oxygen, fermentation Respiration is tightly coupled to other
regenerates the NAD+ needed for pathways 327
glycolysis 311
Plant glycolysis is controlled by its products 312 Respiration in Intact Plants and Tissues 327
Plants respire roughly half of the daily
The Oxidative Pentose Phosphate photosynthetic yield 328
Pathway 312
Respiration operates during photosynthesis 329
The oxidative pentose phosphate pathway
Different tissues and organs respire at different
produces NADPH and biosynthetic
rates 329
intermediates 314
Environmental factors alter respiration rates 329
The oxidative pentose phosphate pathway is
redox-regulated 314 Lipid Metabolism 330
The Citric Acid Cycle 315 Fats and oils store large amounts of energy 331
Mitochondria are semiautonomous Triacylglycerols are stored in oil bodies 331
organelles 315 Polar glycerolipids are the main structural lipids in
Pyruvate enters the mitochondrion and is membranes 332
oxidized via the citric acid cycle 316 Fatty acid biosynthesis consists of cycles of two-
The citric acid cycle of plants has unique carbon addition 334
features 317 Glycerolipids are synthesized in the plastids
and the ER 335
Mitochondrial Electron Transport and
Lipid composition influences membrane
ATP Synthesis 317
function 336
The electron transport chain catalyzes a flow of
Membrane lipids are precursors of important
electrons from NADH to O2 318
signaling compounds 336
The electron transport chain has supplementary
Storage lipids are converted into carbohydrates
branches 320
in germinating seeds 336
ATP synthesis in the mitochondrion is coupled to
electron transport 320 SUMMARY 338
Transporters exchange substrates and
products 322

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XXIV TABLE OF CONTENTS

CHAPTER 12 Assimilation of Mineral Nutrients 343


Nitrogen in the Environment 344 Establishing symbiosis requires an exchange of
Nitrogen passes through several forms in a biogeo- signals 354
chemical cycle 344 Nod factors produced by bacteria act as signals
Unassimilated ammonium or nitrate may be dan- for symbiosis 354
gerous 346 Nodule formation involves phytohormones 355
Nitrate Assimilation 346 The nitrogenase enzyme complex fixes N2 357
Many factors regulate nitrate reductase 347 Amides and ureides are the transported
forms of nitrogen 358
Nitrite reductase converts nitrite to
ammonium 347 Sulfur Assimilation 358
Both roots and shoots assimilate nitrate 348 Sulfate is the absorbed form of sulfur in
plants 358
Ammonium Assimilation 348
Sulfate assimilation requires the reduction of
Converting ammonium to amino acids requires sulfate to cysteine 359
two enzymes 348
Sulfate assimilation occurs mostly in leaves 360
Ammonium can be assimilated via an
alternative pathway 350 Methionine is synthesized from cysteine 360
Transamination reactions transfer nitrogen 350 Phosphate Assimilation 360
Asparagine and glutamine link carbon and
Cation Assimilation 361
nitrogen metabolism 350
Cations form noncovalent bonds with carbon
Amino Acid Biosynthesis 351 compounds 361
Biological Nitrogen Fixation 351 Roots modify the rhizosphere to acquire iron 362
Free-living and symbiotic bacteria fix Iron forms complexes with carbon
nitrogen 351 and phosphate 363
Nitrogen fixation requires anaerobic Oxygen Assimilation 363
conditions 352
The Energetics of Nutrient Assimilation 364
Symbiotic nitrogen fixation occurs in
specialized structures 354 SUMMARY 365

CHAPTER 13 Secondary Metabolites and Plant Defense 369


Secondary Metabolites 370 Some terpenes have roles in growth and
Secondary metabolites defend plants against her- development 373
bivores and pathogens 370 Terpenes defend many plants against
Secondary metabolites are divided into three ma- herbivores 373
jor groups 370 Phenolic Compounds 374
Terpenes 370 Phenylalanine is an intermediate in the
Terpenes are formed by the fusion of five-carbon biosynthesis of most plant phenolics 375
isoprene units 370 Ultraviolet light activates some simple
There are two pathways for terpene phenolics 377
biosynthesis 370 The release of phenolics into the soil may
IPP and its isomer combine to form larger limit the growth of other plants 377
terpenes 371 Lignin is a highly complex phenolic
macromolecule 377

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TABLE OF CONTENTS XXV

There are four major groups of flavonoids 378 Damage by insect herbivores induces
Anthocyanins are colored flavonoids that attract systemic defenses 389
animals 378 Herbivore-induced volatiles have complex
Flavones and flavonols may protect against dam- ecological functions 389
age by ultraviolet light 379 Insects have developed strategies to cope
Isoflavonoids have widespread pharmacological with plant defenses 391
activity 379 Plant Defenses against Pathogens 391
Tannins deter feeding by herbivores 380 Pathogens have developed various strategies to
Nitrogen-Containing Compounds 381 invade host plants 391
Alkaloids have dramatic physiological effects on Some antimicrobial compounds are synthesized
animals 381 before pathogen attack 392
Cyanogenic glycosides release the poison Infection induces additional antipathogen
hydrogen cyanide 384 defenses 392
Glucosinolates release volatile toxins 385 Phytoalexins often increase after pathogen
attack 393
Nonprotein amino acids are toxic to
herbivores 385 Some plants recognize specific pathogen-derived
substances 393
Induced Plant Defenses against Insect Exposure to elicitors induces a signal transduction
Herbivores 386 cascade 394
Plants can recognize specific components of A single encounter with a pathogen may increase
insect saliva 386 resistance to future attacks 394
Jasmonic acid activates many defensive Interactions of plants with nonpathogenic bacteria
responses 387 can trigger induced systemic resistance 395
Some plant proteins inhibit herbivore
digestion 389 SUMMARY 396

UNIT III Growth and Development 401


CHAPTER 14 Signal Transduction 403
Signal Transduction in Plant and Several plant hormone receptors encode
Animal Cells 404 components of the ubiquitination machinery 413
Plants and animals have similar transduction Inactivation of repressor proteins results in a
components 404 gene expression response 414
Receptor kinases can initiate a signal Plants have evolved mechanisms for switching
transduction cascade 406 off or attenuating signaling responses 414
Plants signal transduction components have Cross-regulation allows signal transduction
evolved from both prokaryotic and eukaryotic pathways to be integrated 416
ancestors 406
Signal Transduction in Space and Time 418
Signals are perceived at many locations
within plant cells 408 Plant signal transduction occurs over a wide range
of distances 418
Plant signal transduction often involves
inactivation of repressor proteins 409 The timescale of plant signal transduction ranges
from seconds to years 419
Protein degradation is a common feature in
plant signaling pathways 411 SUMMARY 421

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XXVI TABLE OF CONTENTS

Cell Walls: Structure, Biogenesis,


CHAPTER 15
and Expansion 425
The Structure and Synthesis of Secondary walls form in some cells after
Plant Cell Walls 426 expansion ceases 438
Plant cell walls have varied architecture 426 Patterns of Cell Expansion 441
The primary cell wall is composed of Microfibril orientation influences growth
cellulose microfibrils embedded in a directionality of cells with diffuse growth 441
polysaccharide matrix 428
Cortical microtubules influence the orientation
Cellulose microfibrils are synthesized at of newly deposited microfibrils 443
the plasma membrane 430
Matrix polymers are synthesized in the The Rate of Cell Elongation 443
Golgi apparatus and secreted via vesicles 433 Stress relaxation of the cell wall drives water
Hemicelluloses are matrix polysaccharides uptake and cell elongation 445
that bind to cellulose 433 Acid-induced growth and wall stress relaxation
Pectins are hydrophilic gel-forming components are mediated by expansins 446
of the matrix 434 Many structural changes accompany the
Structural proteins become cross-linked in cessation of wall expansion 448
the wall 437
New primary walls are assembled during SUMMARY 448
cytokinesis 437

CHAPTER 16 Growth and Development 453


Overview of Plant Growth and The differentiation of cortical and endodermal
Development 454 cells involves the intercellular movement of a
Sporophytic development can be divided transcription factor 465
into three major stages 455 Many developmental processes involve the
intercellular movement of macromolecules 467
Embryogenesis: The Origins of Polarity 456
Embryogenesis differs between dicots and Meristematic Tissues: Foundations for
monocots, but also features common Indeterminate Growth 468
fundamental processes 456 The root and shoot apical meristems use similar
Apicalbasal polarity is established early strategies to enable indeterminate growth 469
in embryogenesis 457 The Root Apical Meristem 469
Position-dependent signaling guides The root tip has four developmental zones 469
embryogenesis 458
The origin of different root tissues can be
Auxin may function as a mobile chemical traced to specific initial cells 470
signal during embryogenesis 460
Cell ablation experiments implicate directional
Mutant analysis has helped identify genes signaling processes in determination of cell
essential for embryo organization 461 identity 471
The GNOM protein establishes a Auxin contributes to the formation and
polar distribution of auxin efflux proteins 463 maintenance of the RAM 471
MONOPTEROS encodes a transcription factor Responses to auxin depend on specific
that is activated by auxin 463 transcription factors 472
Radial patterning guides formation of tissue Cytokinin activity in the RAM is required for root
layers 464 development 473

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TABLE OF CONTENTS XXVII

The Shoot Apical Meristem 474 Localized zones of auxin accumulation promote
The shoot apical meristem has distinct zones leaf initiation 480
and layers 474 Spatially regulated gene expression determines
Shoot tissues are derived from several discrete the planar form of the leaf 481
sets of apical initials 475 Distinct mechanisms initiate roots and shoots 483
The locations of PIN proteins influence SAM Senescence and Programmed Cell Death 484
formation 476
Leaf senescence is adaptive and strictly
Embryonic SAM formation requires the regulated 484
coordinated expression of transcription
factors 477 Plants exhibit various types of senescence 485
Negative feedback limits apical meristem size 478 Senescence involves the ordered degradation of
potentially phototoxic chlorophyll 487
Similar mechanisms maintain initials in the RAM
and in the SAM 479 Programmed cell death is a specialized type of
senescence 487
Vegetative Organogenesis 480
SUMMARY 488

Phytochrome and Light Control of Plant


CHAPTER 17
Development 493
The Photochemical and Biochemical Properties Genetic Analysis of Phytochrome
of Phytochrome 494 Function 503
Phytochrome can interconvert between Pr Phytochrome A mediates responses to
and Pfr forms 496 continuous far-red light 504
Pfr is the physiologically active form of Phytochrome B mediates responses
phytochrome 496 to continuous red or white light 504
Characteristics of Phytochrome-Induced Roles for phytochromes C, D, and E
are emerging 504
Responses 497
Phy gene family interactions are complex 504
Phytochrome responses vary in lag time and
escape time 497 PHY gene functions have diversified during
evolution 505
Phytochrome responses can be distinguished by
the amount of light required 497 Phytochrome Signaling Pathways 505
Very lowfluence responses are Phytochrome regulates membrane
nonphotoreversible 497 potentials and ion fluxes 506
Low-fluence responses are photoreversible 498 Phytochrome regulates gene expression 506
High-irradiance responses are proportional to Phytochrome interacting factors (PIFs) act
the irradiance and the duration 499 early in phy signaling 507
Structure and Function of Phytochrome Phytochrome associates with protein kinases
Proteins 499 and phosphatases 507
Phytochrome has several important functional Phytochrome-induced gene expression
domains 500 involves protein degradation 508
Phytochrome is a light-regulated protein Circadian Rhythms 509
kinase 501 The circadian oscillator involves a
Pfr is partitioned between the cytosol and transcriptional negative feedback loop 510
the nucleus 501
Ecological Functions 512
Phytochromes are encoded by a multigene
family 502 Phytochrome enables plant adaptation to
changes in light quality 512

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XXVIII TABLE OF CONTENTS

Decreasing the R:FR ratio causes elongation in Phytochrome responses show ecotypic
sun plants 512 variation 515
Small seeds typically require a high R:FR ratio Phytochrome action can be modulated 515
for germination 513
Reducing shade avoidance responses can SUMMARY 516
improve crop yields 514

Blue-Light Responses: Morphogenesis and


CHAPTER 18
Stomatal Movements 521
The Photophysiology of Blue-Light The Regulation of Blue LightStimulated
Responses 522 Responses 531
Blue light stimulates asymmetric growth and Blue-Light Photoreceptors 532
bending 523
Cryptochromes regulate plant development 532
Blue light rapidly inhibits stem elongation 523
Phototropins mediate blue lightdependent
Blue light stimulates stomatal opening 524 phototropism and chloroplast movements 533
Blue light activates a proton pump at the Zeaxanthin mediates blue-light photoreception
guard cell plasma membrane 527 in guard cells 534
Blue-light responses have characteristic Green light reverses blue lightstimulated
kinetics and lag times 528 opening 536
Blue light regulates the osmotic balance of
guard cells 528 SUMMARY 539
Sucrose is an osmotically active solute in guard
cells 530

Auxin: The First Discovered Plant Growth


CHAPTER 19
Hormone 545
The Emergence of the Auxin Concept 546 Auxin Signal Transduction Pathways 560
The Principal Auxin: Indole-3-Acetic Acid 546 The principal auxin receptors are soluble
protein heterodimers 561
IAA is synthesized in meristems and young
dividing tissues 549 Auxin-induced genes are negatively regulated
by AUX/IAA proteins 561
Multiple pathways exist for the biosynthesis of
IAA 549 Auxin binding to a TIR1/AFB-AUX/IAA
heterodimer stimulates AUX/IAA
Seeds and storage organs contain covalently destruction 562
bound auxin 550
Auxin-induced genes fall into two classes:
IAA is degraded by multiple pathways 550 early and late 562
Auxin Transport 551 Rapid, nontranscriptional auxin responses
Polar transport requires energy and is gravity appear to involve a different receptor
independent 552 protein 562
Chemiosmotic potential drives polar transport 553 Actions of Auxin: Cell Elongation 562
PIN and ABCB transporters regulate cellular auxin Auxins promote growth in stems and coleoptiles,
homeostasis 555 while inhibiting growth in roots 563
Auxin influx and efflux can be chemically The outer tissues of dicot stems are the targets
inhibited 556 of auxin action 563
Auxin transport is regulated by multiple The minimum lag time for auxin-induced
mechanisms 558 elongation is ten minutes 565

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TABLE OF CONTENTS XXIX

Auxin rapidly increases the extensibility of Auxin is redistributed laterally in the root cap 572
the cell wall 565
Developmental Effects of Auxin 573
Auxin-induced proton extrusion increases cell
extension 565 Auxin regulates apical dominance 574
Auxin-induced proton extrusion involves activation Auxin transport regulates floral bud development
and protein mobilization 566 and phyllotaxy 576
Auxin promotes the formation of lateral and
Actions of Auxin: Plant Tropisms 566 adventitious roots 576
Phototropism is mediated by the lateral redistribu- Auxin induces vascular differentiation 576
tion of auxin 566
Auxin delays the onset of leaf abscission 577
Gravitropism involves lateral redistribution of
Auxin promotes fruit development 577
auxin 568
Synthetic auxins have a variety of commercial
Dense plastids serve as gravity sensors 569
uses 578
Gravity sensing may involve pH and calcium ions
(Ca2+) as second messengers 571 SUMMARY 578

Gibberellins: Regulators of Plant Height and


CHAPTER 20
Seed Germination 583
Gibberellins: Their Discovery and Chemical Some enzymes in the GA pathway are
Structure 584 highly regulated 591
Gibberellins were discovered by studying Gibberellin regulates its own metabolism 592
a disease of rice 584 GA biosynthesis occurs at multiple plant organs
Gibberellic acid was first purified from Gibberella and cellular sites 592
culture filtrates 584 Environmental conditions can influence
All gibberellins are based on an ent-gibberellane GA biosynthesis 593
skeleton 585 GA1 and GA4 have intrinsic bioactivity for
stem growth 594
Effects of Gibberellins on Growth and
Development 586 Plant height can be genetically engineered 595
Gibberellins promote seed germination 586 Dwarf mutants often show other phenotypic
defects 595
Gibberellins can stimulate stem and root
growth 586 Auxins can regulate GA biosynthesis 595
Gibberellins regulate the transition from juvenile Gibberellin Signaling: Significance of
to adult phases 587 Response Mutants 596
Gibberellins influence floral initiation and sex GID1 encodes a soluble GA receptor 596
determination 588 DELLA-domain proteins are negative
Gibberellins promote pollen development regulators of GA response 600
and tube growth 588 Mutation of negative regulators of GA may
Gibberellins promote fruit set and produce slender or dwarf phenotypes 600
parthenocarpy 588 Gibberellins signal the degradation of negative
Gibberellins promote early seed regulators of GA response 601
development 588 F-box proteins target DELLA domain proteins
Commercial uses of gibberellins and for degradation 601
GA biosynthesis inhibitors 588 Negative regulators with DELLA domains have
Biosynthesis and Deactivation of agricultural importance 602
Gibberellins 589 Gibberellin Responses: Early Targets
Gibberellins are synthesized via the terpenoid of DELLA Proteins 602
pathway 589

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XXX TABLE OF CONTENTS

DELLA proteins can activate or suppress gene Gibberellin Responses: Anther Development
expression 603 and Male Fertility 607
DELLA proteins regulate transcription by interact- GAMYB regulates male fertility 609
ing with other proteins such as phytochrome- Events downstream of GAMYB in rice aleurone
interacting factors 603 and anthers are quite different 611
Gibberellin Responses: The Cereal Aleurone MicroRNAs regulate MYBs after transcription
Layer 605 in anthers but not in aleurone 611
GA is synthesized in the embryo 605 Gibberellin Responses: Stem Growth 612
Aleurone cells may have two types of GA recep- Gibberellins stimulate cell elongation and
tors 605 cell division 612
Gibberellins enhance the transcription of GAs regulate the transcription of cell cycle
-amylase mRNA 605 kinases 613
GAMYB is a positive regulator of -amylase Reducing GA sensitivity may prevent crop
transcription 607 losses 613
DELLA-domain proteins are rapidly
degraded 607 SUMMARY 614

CHAPTER 21 Cytokinins: Regulators of Cell Division 621


Cell Division and Plant Development 622 Cellular and Molecular Modes of Cytokinin
Differentiated plant cells can resume division 622 Action 629
Diffusible factors control cell division 622 A cytokinin receptor related to bacterial
Plant tissues and organs can be cultured 622 two-component receptors has been
identified 629
The Discovery, Identification, and Properties Cytokinins increase expression of the type-A
of Cytokinins 623 response regulator genes via activation of the
Kinetin was discovered as a breakdown type-B ARR genes 630
product of DNA 623 Histidine phosphotransfer proteins are also
Zeatin was the first natural cytokinin involved in cytokinin signaling 632
discovered 623
The Biological Roles of Cytokinins 632
Some synthetic compounds can mimic
Cytokinins promote shoot growth by increasing
cytokinin action 624
cell proliferation in the shoot apical
Cytokinins occur in both free and bound meristem 632
forms 625
Cytokinins interact with other hormones and with
Some plant pathogenic bacteria, fungi, insects, several key transcription factors 634
and nematodes secrete free cytokinins 625
Cytokinins inhibit root growth by promoting the
Biosynthesis, Metabolism, and Transport exit of cells from the root apical meristem 635
of Cytokinins 625 Cytokinins regulate specific components of
Crown gall cells have acquired a gene for the cell cycle 636
cytokinin synthesis 626 The auxin:cytokinin ratio regulates morphogenesis
IPT catalyzes the first step in cytokinin in cultured tissues 637
biosynthesis 628 Cytokinins modify apical dominance and promote
Cytokinins can act both as long distance and lateral bud growth 638
local signals 628 Cytokinins delay leaf senescence 638
Cytokinins are rapidly metabolized by Cytokinins promote movement of nutrients 639
plant tissues 628

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TABLE OF CONTENTS XXXI

Cytokinins affect light signaling via Cytokinins are involved in the formation of
phytochrome 640 nitrogen-fixing nodules in legumes 641
Cytokinins regulate vascular development 641
SUMMARY 643
Manipulation of cytokinins to alter agriculturally
important traits 641

CHAPTER 22 Ethylene: The Gaseous Hormone 649


Structure, Biosynthesis, and Measurement Ethylene promotes the ripening of some
of Ethylene 650 fruits 659
Regulated biosynthesis determines the Fruits that respond to ethylene exhibit a
physiological activity of ethylene 650 climacteric 659
Ethylene biosynthesis is promoted by several The receptors of never-ripe mutants of tomato
factors 652 fail to bind ethylene 660
Ethylene biosynthesis can be elevated through a Leaf epinasty results when ACC from the root is
stabilization of ACC synthase protein 652 transported to the shoot 660
Various inhibitors can block ethylene Ethylene induces lateral cell expansion 661
biosynthesis 653 There are two distinct phases to growth
inhibition by ethylene 662
Ethylene Signal Transduction Pathways 653
The hooks of dark-grown seedlings are
Ethylene receptors are related to bacterial two-
maintained by ethylene production 662
component system histidine kinases 654
Ethylene breaks seed and bud dormancy in
High-affinity binding of ethylene to its receptor
some species 663
requires a copper cofactor 655
Ethylene promotes the elongation growth of
Unbound ethylene receptors are negative
submerged aquatic species 663
regulators of the response pathway 655
Ethylene induces the formation of roots and
A serine/threonine protein kinase is also involved
root hairs 664
in ethylene signaling 657
Ethylene regulates flowering and sex determination
EIN2 encodes a transmembrane protein 657
in some species 664
Ethylene Regulation of Gene Expression 657 Ethylene enhances the rate of leaf
Specific transcription factors are involved in senescence 664
ethylene-regulated gene expression 657 Ethylene mediates some defense responses 665
Genetic epistasis reveals the order of the Ethylene acts on the abscission layer 665
ethylene signaling components 658 Ethylene has important commercial uses 667
Developmental and Physiological
SUMMARY 668
Effects of Ethylene 659

Abscisic Acid: A Seed Maturation and


CHAPTER 23
Stress-Response Hormone 673
Occurrence, Chemical Structure, and Biosynthesis, Metabolism, and
Measurement of ABA 674 Transport of ABA 674
The chemical structure of ABA determines its ABA is synthesized from a carotenoid
physiological activity 674 intermediate 674
ABA is assayed by biological, physical, and ABA concentrations in tissues are
chemical methods 674 highly variable 676
ABA is translocated in vascular tissue 677

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XXXII TABLE OF CONTENTS

ABA Signal Transduction Pathways 678 ABA regulates seed maturation 684
Receptor candidates include diverse classes of ABA inhibits precocious germination and
proteins 678 vivipary 685
Secondary messengers function in ABA ABA promotes seed storage reserve accumulation
signaling 680 and desiccation tolerance 686
Ca2+-dependent and Ca2+-independent pathways Seed dormancy can be regulated by ABA and
mediate ABA signaling 680 environmental factors 686
ABA-induced lipid metabolism generates second Seed dormancy is controlled by the ratio of ABA
messengers 681 to GA 687
Protein kinases and phosphatases regulate ABA inhibits GA-induced enzyme production 688
important steps in ABA signaling 682 ABA promotes root growth and inhibits shoot
PP2Cs interact directly with the PYR/PYL/RCAR growth at low water potentials 688
family of ABA receptors 683 ABA promotes leaf senescence independently
ABA shares signaling intermediates with other of ethylene 689
hormonal pathways 683 ABA accumulates in dormant buds 689
ABA Regulation of Gene Expression 683 ABA closes stomata in response to water
Gene activation by ABA is mediated by stress 690
transcription factors 684 ABA regulates ion channels and the plasma
membrane ATPase in guard cells 690
Developmental and Physiological Effects
of ABA 684 SUMMARY 693

Brassinosteroids: Regulators of Cell Expansion


CHAPTER 24
and Development 699
Brassinosteroid Structure, Occurrence, Brassinosteroids act locally near their sites of
and Genetic Analysis 700 synthesis 710
BR-deficient mutants are impaired in Brassinosteroids: Effects on Growth and
photomorphogenesis 701 Development 710
The Brassinosteroid Signaling Pathway 703 BRs promote both cell expansion and cell division
BR-insensitive mutants identified the BR cell in shoots 711
surface receptor 703 BRs both promote and inhibit root growth 712
Phosphorylation activates the BRI1 receptor 704 BRs promote xylem differentiation during vascular
BIN2 is a repressor of BR-induced gene development 713
expression 704 BRs are required for the growth of pollen
BES1/BZR1 regulate gene expression 706 tubes 714
BRs promote seed germination 714
Biosynthesis, Metabolism, and Transport of
Brassinosteroids 706 Prospective Uses of Brassinosteroids in
Brassinolide is synthesized from campesterol 707 Agriculture 714
Catabolism and negative feedback contribute to SUMMARY 715
BR homeostasis 708

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TABLE OF CONTENTS XXXIII

CHAPTER 25 The Control of Flowering 719


Floral Meristems and Floral Organ The circadian clock and photoperiodic
Development 720 timekeeping 736
The shoot apical meristem in Arabidopsis changes The coincidence model is based on oscillating
with development 721 light sensitivity 737
The four different types of floral organs are The coincidence of CONSTANS expression
initiated as separate whorls 721 and light promotes flowering in LDPs 737
Two major types of genes regulate floral SDPs use a coincidence mechanism to inhibit
development 722 flowering in long days 739
Meristem identity genes regulate meristem Phytochrome is the primary photoreceptor in
function 722 photoperiodism 739
Homeotic mutations led to the identification of A blue-light photoreceptor regulates flowering
floral organ identity genes 723 in some LDPs 740
Three types of homeotic genes control floral organ Vernalization: Promoting Flowering with
identity 723 Cold 741
The ABC model explains the determination of Vernalization results in competence to flower
floral organ identity 724 at the shoot apical meristem 742
Floral Evocation: Integrating Environmental Vernalization can involve epigenetic changes in
Cues 725 gene expression 742
A range of vernalization pathways may have
The Shoot Apex and Phase Changes 726
evolved 743
Plant development has three phases 726
Juvenile tissues are produced first and are located Long-Distance Signaling Involved in
at the base of the shoot 727 Flowering 744
Phase changes can be influenced by nutrients, The floral stimulus is transported in the
gibberellins, and other signals 728 phloem 744
Competence and determination are two stages in Grafting studies have provided evidence for a
floral evocation 728 transmissible floral stimulus 744

Circadian Rhythms: The Clock Within 730 The Discovery of Florigen 745
Circadian rhythms exhibit characteristic The Arabidopsis protein FLOWERING LOCUS T
features 730 is florigen 746
Phase shifting adjusts circadian rhythms to Gibberellins and ethylene can induce
different daynight cycles 732 flowering 747
Phytochromes and cryptochromes entrain Climate change has already caused measurable
the clock 732 changes in flowering time of wild plants 748
The transition to flowering involves multiple
Photoperiodism: Monitoring Day Length 732 factors and pathways 748
Plants can be classified according to their
photoperiodic responses 732 SUMMARY 749
The leaf is the site of perception of the
photoperiodic signal 734
Plants monitor day length by measuring the length
of the night 734
Night breaks can cancel the effect of the dark
period 735

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XXXIV TABLE OF CONTENTS

CHAPTER 26 Responses and Adaptations to Abiotic Stress 755


Adaptation and Phenotypic Plasticity 756 High Light Stress 764
Adaptations involve genetic modification 756 Photoinhibition by high light leads to the
Phenotypic plasticity allows plants to respond to production of destructive forms of oxygen 764
environmental fluctuations 756 Developmental and Physiological Mechanisms
The Abiotic Environment and its Biological that Protect Plants against Environmental
Impact on Plants 756 Extremes 765
Climate and soil influence plant fitness 757 Plants can modify their life cycles to avoid
Imbalances in abiotic factors have primary and abiotic stress 765
secondary effects on plants 757 Phenotypic changes in leaf structure and behavior
are important stress responses 765
Water Deficit and Flooding 757 The ratio of root-to-shoot growth increases in
Soil water content and the relative humidity of the response to water deficit 769
atmosphere determine the water status of the Plants can regulate stomatal aperture in response
plant 758 to dehydration stress 769
Water deficits cause cell dehydration and Plants adjust osmotically to drying soil by
an inhibition of cell expansion 759 accumulating solutes 769
Flooding, soil compaction, and O2 deficiency are Submerged organs develop aerenchyma tissue in
related stresses 759 response to hypoxia 770
Imbalances in Soil Minerals 760 Plants have evolved two different strategies to
Soil mineral content can result in plant stress protect themselves from toxic ions: exclusion and
in various ways 760 internal tolerance 772
Soil salinity occurs naturally and as the result of Chelation and active transport contribute to
improper water management practices 761 internal tolerance 773
The toxicity of high Na+ and Cl in the cytosol is Many plants have the capacity to acclimate to cold
due to their specific ion effects 761 temperatures 773
Plants survive freezing temperatures by limiting ice
Temperature Stress 762 formation 774
High temperatures are most damaging to The lipid composition of membranes affects their
growing, hydrated tissues 762 response to temperature 775
Temperature stress can result in damaged Plant cells have mechanisms that maintain protein
membranes and enzymes 762 structure during temperature stress 776
Temperature stress can inhibit photosynthesis 763 Scavenging mechanisms detoxify reactive oxygen
Low temperatures above freezing can species 776
cause chilling injury 764 Metabolic shifts enable plants to cope with
Freezing temperatures cause ice crystal formation a variety of abiotic stresses 777
and dehydration 764
SUMMARY 778

APPENDIX ONE A11 GLOSSARY G1


APPENDIX TWO A21 AUTHOR INDEX AI1
APPENDIX THREE A31 SUBJECT INDEX SI1

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