Table Of Content

Marschner’s Mineral Nutrition of Plants Thispageintentionallyleftblank Marschner’s Mineral Nutrition of Plants Fourth Edition Edited by Zed Rengel UWA School of Agriculture and Environment, University of Western Australia, Perth, WA, Australia Ismail Cakmak Faculty of Engineering and Natural Sciences, Sabanci University, Istanbul, Turkey Philip J. White Ecological Sciences, James Hutton Institute, Dundee, Scotland, United Kingdom AcademicPressisanimprintofElsevier 125LondonWall,LondonEC2Y5AS,UnitedKingdom 525BStreet,Suite1650,SanDiego,CA92101,UnitedStates 50HampshireStreet,5thFloor,Cambridge,MA02139,UnitedStates TheBoulevard,LangfordLane,Kidlington,OxfordOX51GB,UnitedKingdom Copyright©2023ElsevierLtd.Allrightsreserved. Nopartofthispublicationmaybereproducedortransmittedinanyformorbyanymeans,electronicormechanical,including photocopying,recording,oranyinformationstorageandretrievalsystem,withoutpermissioninwritingfromthepublisher. 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Tothefullestextentofthelaw,neitherthePublishernortheauthors,contributors,oreditors,assumeanyliabilityforanyinjury and/ordamagetopersonsorpropertyasamatterofproductsliability,negligenceorotherwise,orfromanyuseoroperationof anymethods,products,instructions,orideascontainedinthematerialherein. ISBN:978-0-12-819773-8 ForInformationonallAcademicPresspublications visitourwebsiteathttps://www.elsevier.com/books-and-journals Publisher:NikkiP.Levy AcquisitionsEditor:NancyJ.Maragioglio EditorialProjectManager:EmeraldLi ProductionProjectManager:SelvarajRaviraj CoverDesigner:MatthewLimbert TypesetbyMPSLimited,Chennai,India Contents Listofcontributors xiii 2.5 Factorsinfluencingionuptakebyroots 29 Abouttheeditors xv 2.5.1 Influxtotheapoplasm 29 Foreword xvii 2.5.2 EffectsofpH 30 2.5.3 Metabolicactivity 32 2.5.4 Interactionsamongionsinthe Part I rhizosphere 33 Nutritional physiology 1 2.5.5 Externalconcentration 40 2.5.6 Plantnutritionalstatus 41 2.5.7 Studyingnutritionatconstant 1 Introduction, definition, and tissueconcentration 45 classification of nutrients 3 2.6 Uptakeofionsandwateralongthe ErnestA.Kirkby rootaxis 47 2.7 Radialtransportofionsandwater Summary 3 acrosstheroot 49 1.1 General 3 2.8 Releaseofionsintothexylem 53 1.2 Essentialelementsforplantgrowth 4 2.9 Factorsgoverningionreleaseintothe 1.3 Beneficialelementsforplantgrowth 5 xylemandexudationrate 53 1.4 Anewdefinitionofamineralplant References 56 nutrient 6 1.5 Biochemicalpropertiesandphysiological functionsofnutrientelementsinplants 6 1.6 Variationintheangiospermionome 7 3 Long-distance transport in the References 8 xylem and phloem 73 Furtherreading 9 PhilipJ.WhiteandGuangdaDing Summary 73 2 Ion-uptake mechanisms of 3.1 General 73 individual cells and roots: 3.2 Xylemtransport 74 short-distance transport 11 3.2.1 Compositionofthexylemsap 74 3.2.2 Xylemloading 76 DevrimCoskunandPhilipJ.White 3.2.3 Effectoftranspirationrateon Summary 11 solutetransportinthexylem 82 2.1 General 11 3.2.4 Effectoftranspirationrateon 2.2 Pathwayofsolutesfromtheexternal distributionofelementswithinthe solutionintorootcells 12 shoot 83 2.2.1 Influxtotheapoplasm 12 3.3 Phloemtransport 84 2.2.2 Passageintothecytoplasm 14 3.3.1 Principlesofphloemtransport 2.3 Compositionofbiologicalmembranes 15 andphloemanatomy 84 2.4 Solutetransportacrossmembranes 18 3.3.2 Phloemloadingandthe 2.4.1 Thermodynamicsofsolutetransport 18 compositionofphloemsap 86 2.4.2 Energydemandforsolute 3.3.3 Mobilityinthephloem 88 transport 25 3.3.4 Transferbetweenthexylemand 2.4.3 Thekineticsofsolutetransportin phloem 88 plantroots 25 3.3.5 Phloemunloading 89 v vi Contents 3.4 Relativeimportanceofphloemand 5.2 Relationshipsbetweennutrient xylemforlong-distancetransportof supplyandyield 132 nutrients 89 5.3 Photosyntheticactivityandrelated 3.4.1 General 89 processes 133 3.4.2 Nutrientswithhighphloemmobility 89 5.3.1 Photosyntheticenergyflowand 3.4.3 Nutrientswithlowphloemmobility 90 photophosphorylation 133 3.4.4 Re-translocationandcyclingof 5.3.2 Photoinhibitionand nutrients 91 photooxidation 135 3.5 Remobilizationofnutrients 93 5.3.3 Carbondioxideassimilation 3.5.1 General 93 andphotorespiration 137 3.5.2 Seedgermination 93 5.3.4 C pathwayofphotosynthesis 4 3.5.3 Vegetativestage 93 andCrassulaceanacid 3.5.4 Reproductivestage 94 metabolism 140 3.5.5 Perennials 97 5.3.5 Effectofleafmaturationonits References 97 sink(cid:1)sourcetransition 143 5.3.6 Leafsenescence 144 5.3.7 Feedbackregulationof 4 Uptake and release of elements by photosynthesisbysinkdemand leaves and other aerial plant parts 105 forcarbohydrates 147 ThomasEichertandVictoriaFerna´ndez 5.3.8 Nutritionandphotosynthesis 148 5.4 Photosyntheticarea 151 Summary 105 5.4.1 Individualleafarea 151 4.1 General 105 5.4.2 Leafareaperplant 152 4.2 Uptakeandreleaseofgasesandother 5.4.3 Canopyleafarea(leafarea volatilecompoundsthroughstomata 106 indexandleafareaduration) 153 4.2.1 Volatilenitrogencompounds 106 5.5 Respirationandoxidative 4.2.2 Volatilesulfurcompounds 107 phosphorylation 154 4.3 Uptakeofsolutes 107 5.6 Transportofassimilatesinphloem 4.3.1 General 107 anditsregulation 156 4.3.2 Structureofthecuticle 108 5.6.1 Phloemloadingofassimilates 156 4.3.3 Nutrientuptakethroughthecuticle 109 5.6.2 Mechanismofphloemtransport 4.3.4 Uptakethroughstomata 110 ofassimilates 159 4.3.5 Roleofexternalfactors 111 5.6.3 Phloemunloading 160 4.4 Foliarapplicationofnutrients 114 5.7 Sinkformation 161 4.4.1 General 114 5.7.1 Shootarchitecturefor 4.4.2 Practicalimportanceoffoliar grain/seedyieldformation 162 applicationofnutrients 115 5.7.2 Flowerinitiationand 4.4.3 Foliarfertilizersforpestand development 163 diseasecontrol 118 5.7.3 Pollinationandseed 4.4.4 Foliaruptakeandirrigationmethods 118 development 165 4.5 Leachingofelementsfromleaves 119 5.7.4 Formationofvegetativesink 4.6 Ecologicalimportanceoffoliaruptake organs 167 andleaching 119 5.8 Sinkactivity 168 4.6.1 Foliarleaching 119 5.9 Roleofphytohormonesinthe 4.6.2 Foliarwaterabsorption 120 regulationofthesink(cid:1)source References 121 relationships 170 5.9.1 Structure,sitesofbiosynthesis, 5 Mineral nutrition, yield, and andmaineffectsof (cid:1) source sink relationships 131 phytohormones 171 5.9.2 Phytohormones,signalperception, ErnestKirkby,MiroslavNikolic, andsignaltransduction 175 PhilipJ.White,andGuohuaXu 5.9.3 Effectsofnutritiononthe Summary 131 endogenousconcentrationsof 5.1 General 131 phytohormones 177 Contents vii 5.9.4 Phytohormonesandsinkaction 181 6.5.3 Cellwallstabilization 244 5.10 Sourceandsinklimitationsonyield 183 6.5.4 Cellextensionandsecretory References 185 processes 244 6.5.5 Membranestabilization 245 6.5.6 Cation(cid:1)anionbalanceand 6 Functions of macronutrients 201 osmoregulation 245 6.5.7 Calciumasanintracellular MalcolmJ.Hawkesford,IsmailCakmak, secondmessenger 246 DevrimCoskun,LuitJ.DeKok,HansLambers, 6.5.8 Calciumasasystemicsignal 247 JanK.Schjoerring,andPhilipJ.White 6.5.9 Calciumsupply,plantgrowth, Summary 201 andplantcomposition 248 6.1 Nitrogen 201 6.6 Potassium 249 6.1.1 Nitratetransportinplants 202 6.6.1 General 249 6.1.2 Ammoniumtransportintoand 6.6.2 Compartmentationandcellular withinplants 206 concentrations 250 6.1.3 OrganicNuptake 208 6.6.3 Enzymeactivation 250 6.1.4 Nitrogenassimilation 208 6.6.4 Proteinsynthesis 252 6.1.5 Nitrogensupply,plantgrowth, 6.6.5 Photosynthesis 252 andcomposition 215 6.6.6 Osmoregulation 254 6.1.6 Nitrogen-useefficiency 218 6.6.7 Phloemtransport 257 6.2 Sulfur 219 6.6.8 Energytransfer 258 6.2.1 General 219 6.6.9 Cation(cid:1)anionbalance 258 6.2.2 Sulfateuptake,reduction,and 6.6.10 Stressresistance 258 assimilation 220 6.6.11 Potassiumsupply,plantgrowth, 6.2.3 MetabolicfunctionsofS 221 andplantcomposition 260 6.2.4 Sulfursupply,plantgrowth, References 260 andplantcomposition 224 6.3 Phosphorus 226 6.3.1 General 226 7 Micronutrients 283 6.3.2 Phosphorusasastructural IsmailCakmak,PatrickBrown, element 227 Jose´ M.Colmenero-Flores,SørenHusted, 6.3.3 Roleinenergytransfer 228 BaharY.Kutman,MiroslavNikolic´,ZedRengel, 6.3.4 Compartmentationandregulatory SidselB.Schmidt,andFang-JieZhao roleofinorganicphosphate 229 6.3.5 Phosphorusfractionsandthe Summary 283 roleofphytate 232 7.1 Iron 283 6.3.6 Phosphorussupply,plantgrowth, 7.1.1 General 283 andplantcomposition 234 7.1.2 Iron-containingconstituents 6.4 Magnesium 235 ofredoxsystems 284 6.4.1 General 235 7.1.3 OtherFe-requiringenzymes 287 6.4.2 Bindingform,compartmentation, 7.1.4 Chloroplastdevelopmentand andhomeostasis 235 photosynthesis 287 6.4.3 Chlorophyllandproteinsynthesis 235 7.1.5 Localizationandbindingstate 6.4.4 Enzymeactivation, ofFe 290 phosphorylation,and 7.1.6 RootresponsestoFedeficiency 291 photosynthesis 236 7.1.7 Irondeficiencyandtoxicity 293 6.4.5 Carbohydratepartitioning 238 7.2 Manganese 294 6.4.6 Magnesiumsupply,plantgrowth, 7.2.1 General 294 andcomposition 240 7.2.2 Mn-containingenzymes 294 6.5 Calcium 241 7.2.3 ThefunctionalroleofMnin 6.5.1 General 241 photosynthesis 294 6.5.2 Bindingformand 7.2.4 Manganeseinsuperoxide compartmentation 241 dismutase 296 viii Contents 7.2.5 Manganeseinoxalateoxidase 297 7.8.2 Uptake,transport,and 7.2.6 OtherMn-dependentenzymes 297 homeostasis 350 7.2.7 Proteins,carbohydrates,and 7.8.3 Chargebalance 352 lipids 297 7.8.4 Photosynthesisandchloroplast 7.2.8 Celldivisionandextension 299 performance 352 7.2.9 Manganesedeficiency 299 7.8.5 Cellosmoregulationandturgor 353 7.2.10 Manganesetoxicity 301 7.8.6 Plantwaterbalanceandwater 7.3 Copper 302 relations 354 7.3.1 General 302 7.8.7 Interactionwithnitrate 355 7.3.2 Copperuptakeandtransport 302 7.8.8 Chloridesupply,deficiency, 7.3.3 Copperproteins 303 plantgrowth,andcropyield 356 7.3.4 Carbohydrate,lipid,andN 7.8.9 Chlorinetoxicity 357 metabolism 305 7.8.10 Chlorineasmicro-and 7.3.5 Lignification 306 macronutrient-concluding 7.3.6 Pollenformationandfertilization 307 remarks 358 7.3.7 Copperdeficiencyandtoxicity 308 References 359 7.4 Zinc 310 7.4.1 General 310 8 Beneficial elements 387 7.4.2 Zn-containingenzymes 310 7.4.3 Zn-activatedenzymes 313 JianFengMa,Fang-JieZhao, 7.4.4 Proteinsynthesis 314 ZedRengel,andIsmailCakmak 7.4.5 Carbohydratemetabolism 315 Summary 387 7.4.6 Tryptophanandindoleacetic 8.1 Definition 387 acidsynthesis 316 8.2 Sodium 387 7.4.7 Membraneintegrityandlipid 8.2.1 General 387 peroxidation 317 8.2.2 Essentiality:Naasnutrient 388 7.4.8 Phosphorus-zincinteractions 318 8.2.3 RoleinC species 389 7.4.9 Zincformsandbioavailability 4 8.2.4 SubstitutionofKbyNa 391 ingrains 320 8.2.5 GrowthstimulationbyNa 393 7.4.10 Zincdeficiencyandtoxicity 321 8.2.6 ApplicationofNafertilizers 396 7.5 Nickel 323 8.3 Silicon 397 7.5.1 General 323 8.3.1 General 397 7.5.2 Ni-containingenzymes 324 8.3.2 Uptake,concentration,and 7.5.3 RoleofNiinNmetabolism 325 distribution 397 7.5.4 Nickelconcentrationinplants 327 8.3.3 Beneficialeffects 399 7.5.5 Nickeldeficiencyandtoxicity 327 8.4 Cobalt 402 7.5.6 TolerancetoNitoxicity 328 8.4.1 RoleofCoinplants 402 7.6 Molybdenum 328 8.4.2 Cobaltdeficiencyandtoxicity 403 7.6.1 General 328 8.5 Selenium 405 7.6.2 Molybdenumuptakeandtransport 329 8.5.1 General 405 7.6.3 Nitrogenase 329 8.5.2 Uptakeandtranslocation 405 7.6.4 Nitratereductase 330 8.5.3 Assimilationandmetabolism 407 7.6.5 OtherMo-containingenzymes 331 8.5.4 Beneficialeffectsonplantgrowth 409 7.6.6 Grossmetabolicchanges 333 8.5.5 Biofortification 409 7.6.7 Molybdenumdeficiencyand 8.6 Aluminum 410 toxicity 334 8.7 Otherelements 411 7.7 Boron 336 References 411 7.7.1 General 336 7.7.2 Boroncomplexeswithorganic structures 338 9 Mineral nutrition and crop quality 419 7.7.3 FunctionofB 339 UmitBarisKutman 7.7.4 Borondeficiencyandtoxicity 347 7.8 Chlorine 350 Summary 419 7.8.1 General 350 9.1 Introduction 419 Contents ix 9.2 Technicalquality 420 11 Diagnosis and prediction of 9.2.1 Breadandpasta 420 deficiency and toxicityof nutrients 477 9.2.2 Sugarandoilcrops 421 RichardBell 9.2.3 Fibercrops 422 9.2.4 Processingtomatoes 422 Summary 477 9.2.5 Beerandwine 423 11.1 General 477 9.3 Sensoryquality 423 11.2 Toolsfordiagnosisofnutrient 9.3.1 Effectsofmineralnutritionon disorders 478 visualquality 423 11.2.1 Fieldresponsestonutrient 9.3.2 Effectsofmineralnutritionon supply 478 flavor 425 11.2.2 Diagnosisofnutritional 9.4 Nutritionalquality 426 disordersbyvisiblesymptoms 478 9.4.1 Mineralnutrients,hiddenhunger, 11.2.3 PlantAnalysis 480 andbiofortification 427 11.3 Plantanalysisforprognosisof 9.4.2 Proteinconcentrationandamino nutrientdeficiency 489 acidcomposition 429 11.4 Plantanalysisversussoilanalysis 490 9.4.3 Vitaminsandbioactive References 491 phytochemicals 430 9.5 Shelflifeoffreshfruitsandvegetables 431 9.6 Foodsafety 432 Part II 9.6.1 Toxicelements 432 (cid:1) 9.6.2 HarmfulNcompounds 434 Plant soil relationships 497 9.7 Theyield-qualitydilemma 435 References 436 12 Nutrient availability in soils 499 PetraMarschnerandZedRengel 10 Relationship between mineral nutrition, plant diseases, and pests 445 Summary 499 12.1 General 499 MarkusWeinmann,Kla´raBrada´cˇova´,and 12.2 Chemicalsoilanalysis 499 MiroslavNikolic 12.3 Movementofnutrientstotheroot surface 501 Summary 445 12.3.1 Principlesofcalculations 501 10.1 General 445 12.3.2 Concentrationofnutrients 10.2 Relationshipbetweensusceptibility inthesoilsolution 502 andnutritionalstatusofplants 447 12.3.3 Roleofmassflow 503 10.3 Fungaldiseases 449 12.3.4 Roleofdiffusion 504 10.3.1 Principlesofinfection 449 12.4 Roleofrootdensity 509 10.3.2 RoleofSi 450 12.5 Nutrientavailabilityanddistribution 10.3.3 RoleofNandK 452 ofwaterinsoils 511 10.3.4 RoleofCaandMg 453 12.6 Roleofsoilstructure 513 10.3.5 Roleofphosphateandphosphite 455 12.7 Intensity/quantityratio,plantfactors, 10.3.6 RoleofS 455 andconsequencesforsoiltesting 514 10.3.7 RoleofMn 455 12.7.1 Modelingofnutrientavailability 10.3.8 Roleofothermicronutrients 456 andcropnutrientuptake 515 10.4 Bacterialandviraldiseases 457 References 516 10.4.1 Bacterialdiseases 457 10.4.2 Viraldiseases 457 10.5 Soil-bornefungalandbacterial 13 Genetic and environmental diseases 458 10.6 Pests 461 regulation of root growth and 10.7 Directandindirecteffectsof development 523 fertilizerapplicationonplantsand PengYuandFrankHochholdinger theirpathogensandpests 464 References 467 Summary 523