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FORAGES

THESCIENCE OFGRASSLAND

AGRICULTURE

VOLUMEII

7TH EDITION

FORAGES

THESCIENCE OFGRASSLAND AGRICULTURE

7TH EDITION

DarenD.Redfearn

With93 contributing authors

VOLUMEII

Thisseventheditionfirstpublished2020

©2020JohnWiley&SonsLtd

EditionHistory

©1951,1962,1973,1985,1995IowaStateUniversityPress

©2007BlackwellPublishing

Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmitted,inany formorbyanymeans,electronic,mechanical,photocopying,recordingorotherwise,exceptaspermittedbylaw.Advice onhowtoobtainpermissiontoreusematerialfromthistitleisavailableathttp://www.wiley.com/go/permissions.

TherightofKennethJ.Moore,MichaelCollins,C.JerryNelsonandDarenD.RedfearntobeidentifiedasEditorsof theeditorialmaterialinthisworkhasbeenassertedinaccordancewithlaw.

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HardbackISBN:9781119436577

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PrintedandboundbyCPIGroup(UK)Ltd,Croydon,CR04YY 10987654321

InPraiseofGrass

Nextinimportancetothedivineprofusionofwater,light, andair,thosethreegreatphysicalfactswhichrenderexistencepossible,maybereckonedtheuniversalbeneficence ofgrass.

Grassistheforgivenessofnatureherconstant benediction....Forestsdecay,harvestsperish,flowersvanish,butgrassisimmortal.Ityieldsnofruitin earthorair,andyetshoulditsharvestfailinasingleyear, faminewoulddepopulatetheearth.

Grasssoftenstherudeoutlineoftheworld.Itstenaciousfibersholdtheearthinplace.Itinvadesthesolitude ofdeserts,climbstheinaccessibleslopesandforbiddingpinnaclesofmountains,modifiesclimates,and determinesthehistory,character,anddestinyofnations.

JohnJamesIngalls KansasMagazine 1872

GrasslandScience

Whoevercouldmake...twobladesofgrasstogrowuponaspotofgroundwhereonlyone grewbefore,woulddeservebetterofmankind,anddomoreessentialservicetohiscountry thanthewholeraceofpoliticiansputtogether.

JonathanSwift fromGulliver’stravels,1726

Contents

Preface,xiii

ListofContributors,xv

Dedication,xxi

PARTI FORAGEPLANTS,1

1Perspectives,Terminology,andClassification,3

C.JerryNelson,ProfessorEmeritus,KennethJ.Moore,DistinguishedProfessor,MichaelCollins,ProfessorEmeritusand DarenD.Redfearn,AssociateProfessor

2GrassMorphology,23

C.JerryNelson,ProfessorEmeritusandKennethJ.Moore,DistinguishedProfessor

3LegumeStructureandMorphology,51

JohnJennings,ProfessorandJamieFoster,Professor

4CarbonMetabolisminForagePlants,65

JeffreyJ.Volenec,ProfessorandC.JerryNelson,ProfessorEmeritus

5MineralNutrientAcquisitionandMetabolism,85

SylvieM.Brouder,WickershamChairandProfessorandJeffreyJ.Volenec,Professor

6Plant-WaterRelationsinForageCrops,113

JenniferW.MacAdam,ProfessorandC.JerryNelson,ProfessorEmeritus

7GrowthandDevelopment,127

RobertB.Mitchell,ResearchAgronomist,DarenD.Redfearn,AssociateProfessorandKennethJ.Moore, DistinguishedProfessor

PARTII FORAGEECOLOGY,149

8Climate,Climate-ChangeandForageAdaptation,151

VernS.Baron,ResearchScientistandGillesBélanger,ResearchScientist

9PlantInteractions,187

JohnA.Guretzky,GrasslandSystemsEcologist

10Plant-HerbivoreInteractions,201

LynnE.Sollenberger,DistinguishedProfessorandMarceloO.Wallau,AssociateProfessor

11NutrientCyclinginForageProductionSystems,215

DavidA.Wedin,ProfessorandMichaelP.Russelle,SoilScientist(Retired)

12ForagesforConservationandImprovedSoilQuality,227

JohnF.Obrycki,ORISEFellowandDouglasL.Karlen,SoilScientist(Retired)

13ForagesandtheEnvironment,249

MattA.Sanderson,ResearchAgronomistandResearchLeader(Retired)andMarkA.Liebig,SoilScientist

PARTIII FORAGESPECIES,261

14Cool-SeasonLegumesforHumidAreas,263

CraigC.Sheaffer,Professor,GeraldW.Evers,ProfessorEmeritusandJacobM.Jungers,AssociateProfessor

15LegumesforTropicalandSubtropicalAreas,277

WilliamD.Pitman,ProfessorandJoãoM.B.Vendramini,AssociateProfessor

16Cool-SeasonGrassesforHumidAreas,297

MichaelD.Casler,ResearchGeneticist,RobertL.Kallenbach,AssociateDeanandGeoffreyE.Brink,Research Agronomist

17GrassesforAridandSemiaridAreas,313

DarenD.Redfearn,AssociateProfessor,KeithR.Harmoney,RangeScientistandAlexanderJ.Smart,Professorand RangelandManagementSpecialist

18Warm-SeasonGrassesforHumidAreas,331

LynnE.Sollenberger,DistinguishedProfessor,JoãoM.B.Vendramini,AssociateProfessor,CarlosG.S.Pedreira, AssociateProfessorandEstebanF.Rios

19ForbsandBrowseSpecies,347

DavidP.Belesky,ClinicalAssociate&Director,JohnW.Walker,ProfessorandResidentDirector,KimberlyA.Cassida, ForageExtensionSpecialistandJamesP.Muir,Professor

PARTIV FORAGESYSTEMS,367

20SystemsforTemperateHumidAreas,369

JeromeH.Cherney,Professor,RobertL.Kallenbach,AssociateDeanandValentínD.PicassoRisso,AssistantProfessor

21ForageSystemsfortheTemperateSubhumidandSemiaridAreas,387

JohnR.Hendrickson,ResearchRangelandManagementSpecialistandCoreyMoffet,ResearchRangelandManagement Specialist

22SystemsfortheWarmHumidAreas,407

WilliamD.Pitman,ProfessorandMontgomeryW.Alison,ExtensionForageSpecialist

23SystemsforHumidTransitionAreas,419

RenataN.Oakes,AssistantProfessorandDennisW.Hancock,CenterDirector

24ForageSystemsforAridAreas,433

DanielH.Putnam,ForageExtensionSpecialistandTimDelCurto,ProfessorandNancyCameronChair

PARTV FORAGEPRODUCTIONANDMANAGEMENT,453

25ForageEstablishmentandRenovation,455

MarvinH.Hall,Professor,YoanaC.Newman,AssociateProfessorandJessicaA.Williamson,AssistantProfessor

26FertilizationandNutrientManagement,473

DavidJ.Barker,ProfessorandStevenW.Culman,Professor

27IrrigationandWaterManagement,497

L.NielAllen,AssociateProfessorandIrrigationSpecialistandJenniferW.MacAdam,ProfessorofPlants, SoilsandClimate

28WeedManagement,515

RobertA.Masters,RangelandScientist(Retired),ByronB.Sleugh,ForageAgronomistandE.ScottFlynn, ForageAgronomist

29InsectManagement,535

R.MarkSulc,Professor,WilliamO.Lamp,ProfessorandG.DavidBuntin,Professor

PARTVI FORAGEIMPROVEMENT,551

30ForageBreeding,553

MichaelD.Casler,ResearchGeneticistandKennethP.Vogel,ResearchGeneticist(retired)

31BiotechnologyandMolecularApproachestoForageImprovement,567

E.CharlesBrummer,ProfessorandZeng-YuWang,Professor

32SeedProduction,581

JeffreyJ.Steiner,AssociateDirectorandTimL.Springer,ResearchAgronomist

PARTVII FORAGEQUALITY,593

33CarbohydrateandProteinNutritionalChemistryofForages,595

RonaldD.Hatfield,ResearchPlantPhysiologistandKennethF.Kalscheur,ResearchDairyScientist

34DigestibilityandIntake,609

DavidR.Mertens,PresidentandResearchDairyScientist(Retired)andRichardJ.Grant,Presidentand ResearchScientist

35PlantChemistryandAntiqualityComponentsinForage,633

NicholasS.Hill,ProfessorandCraigA.Roberts,Professor

36LaboratoryMethodsforEvaluatingForageQuality,659

WilliamP.Weiss,ProfessorandMaryBethHall,ResearchAnimalScientist

37AnimalMethodsforEvaluatingForageQuality,673

EricS.Vanzant,AssociateProfessor,RobertC.Cochran,ProfessorandWayneK.Coblentz,ResearchDairy Scientist/Agronomist

38PredictingForageQuality,687

DebbieJ.Cherney,ProfessorandDavidParsons,Professor

39FactorsAffectingForageQuality,701

KennethJ.Moore,CharlesF.Curtiss,DistinguishedProfessor,AndrewW.Lenssen,ProfessorandStevenL.Fales, EmeritusProfessor

PARTVIII FORAGEHARVESTINGANDUTILIZATION,719

40Post-HarvestPhysiology,721

WayneK.Coblentz,ResearchDairyScientist/Agronomist

41HayHarvestandStorage,749

C.AlanRotz,AgriculturalEngineer,KevinJ.Shinners,ProfessorandMatthewDigman,AssistantProfessor

42SilageProduction,767

RichardE.Muck,AgriculturalEngineer,LiminKungJr.,ProfessorandMichaelCollins,ProfessorandDirector Emeritus

43Biomass,Energy,andIndustrialUsesofForages,789

MattA.Sanderson,ResearchAgronomistandResearchLeader(Retired),PaulAdler,ResearchAgronomistandNealP. Martin,Director(Retired)

PARTIX PASTUREMANAGEMENT,801

44PastureDesignandGrazingManagement,803

LynnE.Sollenberger,DistinguishedProfessor,YoanaC.Newman,AssociateProfessorandBisoondatMacoon,Research Professor

45GrazingAnimalNutrition,815

GregoryLardy,DepartmentHeadandRichardWaterman,ResearchAnimalScientist

46GrazingAnimalBehavior,827

KarenL.Launchbaugh,HeadyProfessor

47Forage-InducedAnimalDisorders,839

TimA.McAllister,PrincipalResearchScientist,GabrielRibeiro,AssistantProfessor,KimStanford,ResearchScientist andYuxiWang,ResearchScientist

48GrazingSystemsandStrategies,861

MichaelCollins,ProfessorandDirectorEmeritus,KennethJ.Moore,DistinguishedProfessor,C.JerryNelson,Professor EmeritusandDarenD.Redfearn,AssociateProfessor

CommonandScientificNamesofForages,883

Glossary,893

Index,919

Preface

ForagesinTransition

Itisdauntingtoconsiderhowtoincreasethefoodsupply whileconservingnaturalresourcestofeedtheexpected10 billionpeopleworldwideby2050.Thismustoccurwith lessland,lesswater,lessfossilfuel,highercostsoflabor, andwillrequiremoreefficientuseofinputs.And,itmust bedonewhileprotectingtheenvironmentinthefaceof globalclimatechangeandgreaterpublicdemandforsustainability.Foragesandpastureswillplayacriticalroleby effectivelyusinglowerqualitylandresources,whilesimultaneously,supplyinganadequatequantityofhigh-quality andsafeproducts,especiallyanimalproducts.Emphasis willincreaseforforagesandpasturestocontributespecific ecosystemservices.

MuchofthelandresourceofNorthAmericaisoccupiedbygrasslandsandforagesmanagedbyranchersand farmersforyield,qualityandpersistence.However,the socialclimatesurroundingagricultureisrapidlychanging asthepublicbecomesmoreconcernedandevendistrustfulaboutmotivesandprioritiesoflandmanagement forincomeoversustainability.Howwillresearchand technicaladvancementofforagesandpasturesaddress thenon-productionfactorswhilemovingthediscipline forward?

VolumeIofthe7th editionofForages,anIntroduction toGrasslandAgriculture(2018),servesprimarilyasan undergraduatetextbook.Itemphasizesbasicrolesof thediversearrayofforageplants,theiradaptation,and principlesofmanagementpracticesusedforefficient animalproductionthatissustainable.VolumeIIof the7th editionofForages,theScienceofGrassland Agriculture(2020)givesmoredetailonhowbiological andphysicalprocessesincellsandtissuesaffectgrowth, foragequalityandpersistenceofindividualplants.We thenintegratethebasicknowledgeaboutindividual plantstotheirinteractioninplantcommunities,whether

harvestedmechanicallyorgrazedbyanimals,andhow theycontributeecosystemservices.

Forageyieldinresearchplotshasincreasedvery littleoverthepasthalfcentury.Relativefocusischangingfromincreasingyieldtoreducinginputcostsand improvingandretainingforagequality.Newcultivars andstrategiesfordiseaseandinsectcontrolhelpprotect yieldandimprovebothanimalperformanceandstand persistence.Newharvestingequipmentimprovesleaf retentionandshortensdryingtimetoreduceweathering losses.Improvedbalewrappingandsilagepreservation technologiesfurtherhelpretaindigestiblecomponents. Globalpositioninganddroneswillfindimportantuses forprecisionfarmingtoincreasemanagementefficiency.

Atthesametime,thepublicdesiresincreasedemphasisonecology,climatechange,ecosystemservices,animal welfare,andsustainableforageandpasturemanagement. Theseconcernshaveledtostrongerlinksamongforage scientists,animalscientists,ecologists,climatologistsand socialscientiststoformtransdisciplinaryfoundationsfor managingforagesandpastures.Thebroaderroleofforages andpastureswillleadtonewpoliciestoprovidequalityanimalproductsaswellasvaluableecosystemservices. Newsciencewillestablishthebestpoliciesandpractices.

ForagesNeedInnovation

Foragesandpasturescaneffectivelyuselandresources thatdonotcompetedirectlywithgrainandoilseed cropproduction.Ruminantsarecriticalsincetheyhave naturaladvantagesinconvertingfibrousplantmaterial intohighnutritionalvaluemeatandmilkproducts. Hundredsofplantscouldbecomesignificantforages inspecificenvironments,andbiotechnologywillhelp improvespeciesalreadyused.Directuseofperennialsfor renewableenergysourcescanreducedependenceonfossil fuels.Forageswillbemoreintegralcomponentsofcrop rotations,covercropsandvegetativewaterwaysforfeed

sourcesanderosioncontrol.Perenniallegumesincrop rotationswillprotectthesoilandsupportwildlifewhile providingfixednitrogenforsubsequentcrops.

Fortunately,therearemanynewtechnologiesinthe pipelinesuchasglobalpositioningsystems,precision agriculture,drones,improvedharvestingandpackaging machinery,saferpesticides,improvedefficiencyoffertilizeruseandmanyfindingsfrombiotechnologythatare leadingtomajorchangesinplantandanimalagriculture. Scientistsarelearningaboutmanagingmarginalsoils, howecosystemswork,hownewtechnologiesmight betransferrabletootherareasandhowthebenefitsof plantdiversityassistinmaintainingecosystemservices. Theprivatesectorwillcontinuetohelpbydeveloping newcultivars,improvedfarmmachinery,newresearch methodologiesandinstrumentsformonitoringhayfields, pasturesandanimalbehavior.

ForagesandtheRoleofVolumeII

ForVolumeIIofForages,TheScienceofGrasslandAgriculture,authorsassembledathoroughreviewofrelevant literaturetoglean,evaluateandintegratethemostimportantfactorsforcurrentandpotentialuse.Unfortunately, thenumberofforageresearchersintheUSandCanada isdecreasing,similartotrendsinEurope,Australia,New ZealandandSouthAmerica.Thisrequiresmoreuseof internationalliteraturewhentheinformationistransferableorisvalidatedormodifiedinthenewenvironment. Inaddition,especiallyatbasiclevels,thereisaneedtouse dataandevaluationsfromnon-foragespeciestoprovide insighttoimportantfeaturesofforageandpastureplants. Moretransdisciplinaryresearchwithsocialandenvironmentalscientistshasaidedevaluationofapplicationsfor economicviabilityandsocialacceptance.

Asafirstpriority,authorsconsideredhowresearch improvesadaptation,qualityandpersistenceofforageand pastureplants.Second,authorsevaluatedtechnologies andmanagementsystemsforsustainabilitywithinafield orpasture.Insystemschapters,theyconsideredforages

andpasturesascomponentswhenscaledtocropping orlivestocksystemswithinalargerarea.Third,authors consideredpotentialeffectsofresourcelimitationsand pendingclimatechangetosupportproductionandprovideecosystemservices.Collectively,VolumeIIpresents acomprehensiveassessmentofforagesandtheirrolesin agriculturalsystemsthatarechangingincharacterand function.

ThankstoContributors

Theeditorsareveryappreciativeofthecontributionsof the93authorswhodeliveredthisworkthroughtheir vision,commitmentandknowledge.Theirgenerosity, goodwillandtalentmadethis7th editionofVolume IIpossible.Thecompletededitionalsocontinuesthe traditionofprovidingthemostcomprehensivereference bookavailableonforagesandgrasslandsthatiswritten bynationalleadersintheirareasofeducation,extension, andresearchexpertise.

Insomechapters,conceptsanddescriptionsinclude materialfromchaptersonsimilartopicsinearliereditions, especiallythe5th and6th editions.Thecurrentauthorsand editorsareindebtedtothoseauthorswhohelpedform thefoundationandformatforchaptersinthe7th edition. Withgreatrespect,wethankthoseearlierauthorsfortheir effortstoadvancethescienceofgrasslandagricultureand therolesofforagesandpasturesindynamicecosystems.

KenMooreprovidedadministrativeleadershipforthe projectandalsoeditedandco-authoredchapters.Michael Collins,JerryNelson,andDarenRedfearnsharedinthe editorialworkandalsoco-authoredchapters.Wehope youcanlearnfromandbereassuredandstimulatedby thepublication.Wewelcomeyourresponsesaboutour collectiveeffort,bothnegativeandpositive.

KennethJ.Moore

MichaelCollins C.JerryNelson DarenD.Redfearn

ListofContributors

PaulAdler ResearchAgronomist, PastureSystemsandWatershedManagementResearch Unit, USDA-AgriculturalResearchService, UniversityPark,PA,USA

MontgomeryW.Alison ExtensionForageSpecialist, LouisianaStateUniversityAgriculturalCenter, Winnsboro,LA,USA

L.NielAllen AssociateProfessorandIrrigationSpecialist, UtahStateUniversity, Logan,UT,USA

DavidJ.Barker ProfessorofHorticultureandCropScience, TheOhioStateUniversity, Columbus,OH,USA

VernS.Baron ResearchScientist, AgricultureandAgri-FoodCanada, Lacombe,AB,Canada

GillesBélanger ResearchScientist, AgricultureandAgri-FoodCanada, Sainte-Foy,PQ,Canada

DavidP.Belesky ClinicalAssociateProfessor&DirectorDavisCollege FarmSystem, WestVirginiaUniversity, Morgantown,WV,USA

GeoffreyE.Brink ResearchAgronomist,USDA-AgriculturalResearch Service, USDairyForageResearchCenter, Madison,WI,USA

SylvieM.Brouder WickershamChairofExcellenceinAgricultural ResearchandProfessorofAgronomy, PurdueUniversity, WestLafayette,IN,USA

E.CharlesBrummer Professor, UniversityofCalifornia, Davis,CA,USA

G.DavidBuntin ProfessorofEntomology, UniversityofGeorgia, Griffin,GA,USA

MichaelD.Casler USDA-AgriculturalResearchService,USDairyForage ResearchCenter Madison,WI,USA

KimberlyA.Cassida ForageExtensionSpecialist,MichiganStateUniversity EastLansing,MI,USA

DebbieJ.Cherney ProfessorofAnimalScience, CornellUniversity, Ithaca,NY,USA

JeromeH.Cherney ProfessorofSoilandCropSciences, CornellUniversity, Ithaca,NY,USA

WayneK.Coblentz InstituteforEnvironmentallyIntegratedDairy Management, USDairyForageResearchCenter, Marshfield,WI,USA

RobertC.Cochran Professor,KansasStateUniversity, Manhattan,KS,USA

MichaelCollins ProfessorandDirectorEmeritus,DivisionofPlant Sciences, UniversityofMissouri, Manchester,KY,USA

StevenW.Culman Professor,SchoolofEnvironmentandNaturalResources, TheOhioStateUniversity, Columbus,OH,USA

TimDelCurto ProfessorandNancyCameronChair, MontanaStateUniversity, Bozeman,MT,USA

MatthewDigman AssistantProfessorofAgriculturalEngineering, UniversityofWisconsin, RiverFalls,WI,USA

GeraldW.Evers ProfessorEmeritusofSoilandCropSciences, TexasA&MUniversity, Overton,TX,USA

StevenL.Fales EmeritusProfessorofAgronomy, IowaStateUniversity, Ames,IA,USA

E.ScottFlynn Agronomist,CortevaAgriscience, LeesSummit,MO,USA

JamieFoster AssociateProfessorofForageAgronomy,TexasA&M AgriLifeResearch, Beeville,TX,USA

JohnA.Guretzky GrasslandSystemsEcologist, UniversityofNebraska, Lincoln,NE,USA

RichardJ.Grant PresidentandResearchScientist, TheWilliamH.MinerAgriculturalResearchInstitute, Chazy,NY,USA

MarvinH.Hall ProfessorofCropandSoilSciences, PennsylvaniaStateUniversity UniversityPark,PA,USA

MaryBethHall ResearchAnimalScientist,USDA-AgriculturalResearch Service, USDairyForageResearchCenter Madison,WI,USA

DennisW.Hancock CenterDirector,USDA-AgriculturalResearchService, USDairyForageResearchCenter, Madison,WI,USA

KeithR.Harmoney RangeScientist,KansasStateUniversity, Hays,KS,USA

RonaldD.Hatfield ResearchPlantPhysiologist,USDA-Agricultural ResearchService, USDairyForageResearchCenter, Madison,WI,USA

JohnR.Hendrickson ResearchRangelandManagementSpecialist, USDA-AgriculturalResearchService Mandan,ND,USA

NicholasS.Hill ProfessorofCropandSoilSciences, TheUniversityofGeorgia, Athens,GA,USA

JohnJennings

ProfessorofAnimalScience-Forages, UniversityofArkansas, LittleRock,AR,USA

JacobM.Jungers

AssistantProfessorofAgronomyandPlantGenetics, UniversityofMinnesota, St.Paul,MN,USA

RobertL.Kallenbach AssociateDean,AgricultureandEnvironmentExtension, UniversityofMissouri, Columbia,MO,USA

KennethF.Kalscheur ResearchDairyScientist,USDA-AgriculturalResearch Service, USDairyForageResearchCenter, Madison,WI,USA

DouglasL.Karlen SoilScientist(Retired), USDA-AgriculturalResearchService, Ames,IA,USA

LiminKung,Jr. ProfessorofAnimalandFoodSciences, UniversityofDelaware, Newark,DE,USA

WilliamO.Lamp ProfessorofEntomology, UniversityofMaryland, CollegePark,MD,USA

GregoryLardy DepartmentHead,AnimalSciences, NorthDakotaStateUniversity, Fargo,ND,USA

KarenL.Launchbaugh HeadyProfessorofRangelandEcology, UniversityofIdaho, Moscow,ID,USA

AndrewW.Lenssen ProfessorofAgronomy, IowaStateUniversity, Ames,IA,USA

MarkA.Liebig SoilScientist,USDA-AgriculturalResearchService,

NorthernGreatPlainsResearchLaboratory, Mandan,ND,USA

JenniferW.MacAdam ProfessorofPlants,SoilsandClimate, UtahStateUniversity, Logan,UT,USA

BisoondatMacoon ResearchProfessor, MississippiStateUniversity, Raymond,MS,USA

NealP.Martin Director(Retired),USDA-AgriculturalResearch Service, USDairyForageResearchCenter, Madison,WI,USA

RobertA.Masters RangelandScientist(Retired), CortevaAgriscience, Indianapolis,IN,USA

TimA.McAllister PrincipalResearchScientist,AgriculturalandAgri-Food Canada, Lethbridge,AB,Canada

DavidR.Mertens President,MertensInnovation&ResearchLLC, Belleville,WI,USAand ResearchDairyScientist(Retired),USDA-Agricultural ResearchService, USDairyForageResearchCenter, Madison,WI,USA

RobertB.Mitchell ResearchAgronomist,USDA-AgriculturalResearch Service, Lincoln,NE,USA

CoreyMoffet ResearchRangelandManagementSpecialist, USDA-AgriculturalResearchService, Woodward,OK,USA

KennethJ.Moore

CharlesF.CurtissDistinguishedProfessorinAgriculture andLifeSciencesandPioneerHi-BredProfessorof Agronomy, IowaStateUniversity, Ames,IA,USA

RichardE.Muck AgriculturalEngineer,USDA-AgriculturalResearch Service(Retired), USDairyForageResearchCenter, Madison,WI,USA

JamesP.Muir ProfessorGrasslandEcology, TexasA&MAgriLifeResearch&ExtensionCenter, Stephenville,TX,USA

C.JerryNelson ProfessorEmeritusofPlantSciences, UniversityofMissouri, Columbia,MO,USA

YoanaC.Newman AssociateProfessorofPlantandEarthScience, UniversityofWisconsin, RiverFalls,WI,USA

RenataN.Oakes AssistantProfessorofForageSystemsandManagement, UniversityofTennessee, SpringHill,TN,USA

JohnF.Obrycki ORISEFellow,USDANationalLaboratoryfor AgricultureandtheEnvironment, Ames,IA,USA

DavidParsons ProfessorofCropScience, SwedishUniversityofAgriculturalSciences(SLU) Umeå,Sweden

CarlosG.S.Pedreira AssociateProfessorofAnimalScience, UniversityofSãoPaulo, SãoPaulo,Brazil

ValentínD.PicassoRisso AssistantProfessorofAgronomy, UniversityofWisconsin, Madison,WI,USA

WilliamD.Pitman Professor, LouisianaStateUniversityAgriculturalCenter, Homer,LA,USA

DanielH.Putnam ForageExtensionSpecialist, UniversityofCalifornia, Davis,CA,USA

DarenD.Redfearn AssociateProfessorofAgronomy, UniversityofNebraska, Lincoln,NE,USA

GabrielRibeiro AssistantProfessor AnimalandPoultryScience, UniversityofSaskatchewan, Saskatoon,SK,Canada

EstebanF.Rios AssistantProfessorofAgronomy, UniversityofFlorida, Gainesville,FL,USA

CraigA.Roberts ProfessorofAgronomy, UniversityofMissouri, Columbia,MO,USA

C.AlanRotz AgriculturalEngineer,USDA-AgriculturalResearch Service, USDairyForageResearchCenter, Madison,WI,USA

MichaelP.Russelle SoilScientist(Retired),USDA-AgriculturalResearch Service, St.Paul,MN,USA

MattA.Sanderson ResearchAgronomistandResearchLeader(Retired), USDA-AgriculturalResearchService, StateCollege,PA,USA

CraigC.Sheaffer ProfessorofAgronomyandPlantGenetics, UniversityofMinnesota, St.Paul,MN,USA

KevinJ.Shinners ProfessorofAgriculturalEngineering, UniversityofWisconsin, Madison,WI,USA

ByronB.Sleugh Agronomist,CortevaAgriscience, Indianapolis,IN,USA

AlexanderJ.Smart ProfessorandRangelandManagementSpecialist, SouthDakotaStateUniversity, Brookings,SD,USA

LynnE.Sollenberger DistinguishedProfessorofAgronomy, UniversityofFlorida, Gainesville,FL,USA

TimL.Springer ResearchAgronomist,USDA-AgriculturalResearch Service, Woodward,OK,USA

KimStanford ResearchScientist,AlbertaAgricultureandForestry, Lethbridge,AB,Canada

JeffreyJ.Steiner AssociateDirector,GlobalHempInnovationCenter, OregonStateUniversity, Corvalis,OR,USA

R.MarkSulc ProfessorofHorticultureandCropScience, TheOhioStateUniversity, Columbus,OH,USA

EricS.Vanzant AssociateProfessor,UniversityofKentucky, Lexington,KY,USA

JoãoM.B.Vendramini

AssociateProfessorofAgronomy,RangeCattleResearch andEducationCenter, UniversityofFlorida, Ona,FL,USA

KennethP.Vogel USDA-AgriculturalResearchService(retired), Lincoln,NE,USA

JeffreyJ.Volenec ProfessorofAgronomy,PurdueUniversity, WestLafayette,IN,USA

JohnW.Walker ProfessorandResidentDirectorofResearch,Texas A&MAgriLifeResearchandExtensionCenter, SanAngelo,TX,USA

YuxiWang ResearchScientist,AgricultureandAgri-FoodCanada, Lethbridge,AB,Canada

Zeng-YuWang Professor, QingdaoAgriculturalUniversity, Yantai,China

MarceloO.Wallau AssistantProfessorofAgronomy, UniversityofFlorida, Gainesville,FL,USA

RichardWaterman ResearchAnimalScientist,USDA-Agricultural ResearchService,FortKeoghLivestockandRange ResearchLaboratory, MilesCity,MT,USA

DavidA.Wedin Professor, SchoolofNaturalResources, UniversityofNebraska, Lincoln,NE,USA

WilliamP.Weiss ProfessorofAnimalSciences, OhioAgriculturalResearchandDevelopmentCenter, TheOhioStateUniversity, Wooster,OH,USA

JessicaA.Williamson AssistantProfessorofCropandSoilSciences, PennsylvaniaStateUniversity, UniversityPark,PA,USA

Dedication

ThisvolumeisdedicatedtothememoryofDrs.Steven LouisFales,LowellE.MoserandWalterF.Wedin. Devotedandpassionategrasslandersall,theywerealso highlyproductiveresearchersandenthusiasticeducators. Theyinspiredandtrainedmanyoftheauthorscontributingtothisvolume.Theirlivesandcareerscrossedpaths manytimesovertheyearsandallthreewerecontributors toearliereditionsofForages.

Lowelleditedseveralimportantbooksandmonographs relatedtoforagesincluding Cool-SeasonForageGrasses and Warm-Season(C4)Grasses,bothpublishedby theTri-Societies(ASA-CSSA-SSSA).WaltandSteve co-edited Grassland:QuietnessandStrengthforaNew AmericanAgriculture theirhomageto Grass,the1948 YearbookofAgriculture

Thisvolumeisalsorespectfullydedicated1 : TotheMemoryofThose goneonbefore,who,envisioningtheneedsofthefutureandthepossibilityofbetterthings,livedpurposively,givingofthemselves. InRecognitionofThose ofourownday,who,endowed withleadershipabilityinresearchandeducation, continuetostimulateustomoreproductiveeffort. FortheInspirationofThose whotodayfollowon,but whotomorrow,buildinguponestablishedfoundations,willbechargedwiththeresponsibilityofsolvingproblemswithwhichthoseoftheirdaywillbe confronted.

1 From,Hughes,H.D.,Heath,M.E.,andMetcalfe,D.S.(eds.) (1951). Forages:TheScienceofGrasslandAgriculture ,1e.Ames, IA:TheIowaStateCollegePress.

StevenL.Fales
LowelE.Moser
WalterF.Wedin

PART I FORAGEPLANTS

Amixedstandofalfalfaandtimothy.TimothymixtureswithalfalfainKentuckyprovidemixedforageonthefirstcuttingor grazingbutnearlypurealfalfathroughtheremainderofthegrowingseason. Source: PhotocourtesyofMikeCollins.

PartIcoversbasicphysiologicandphysicalpropertiesof foragespeciesatthecellularandwhole-plantlevelsthat guidegeneticimprovementandunderscoremanagement practices.Thegoalsaretoimproveyieldandqualityofthe biomassandresistancetobioticandabioticstresses.These processesoftenhavenegativecorrelationsthatarespecies dependent,andresponsesofspaced-plantsmaynotreflect

Forages:TheScienceofGrasslandAgriculture,VolumeII,SeventhEdition. EditedbyKennethJ.Moore,MichaelCollins,C.JerryNelsonandDarenD.Redfearn. ©2020JohnWiley&SonsLtd.Published2020byJohnWiley&SonsLtd. theirpropertieswhengrownindensestandsormixtures. Criticaltopicssuchasphotosynthesis,rootgrowth, canopyarchitecture,lignificationofcellwallsandpresenceofantiqualityfactorssuchasalkaloidsinleavesneed tocontinuetobeevaluated.Mostperennialforageplants arepolyploidsandcross-pollinated,makingitdifficult toidentifyandtransfergenesusingbiotechnology,but

CRISPR-Cas9andothernewtechnologiesareopening newwaystosupplementtraditionalbreedingmethods. Geneticpotentialforgrowthandpersistenceset theupperlimitsforyield.Managementstrategiesutilize resourcesefficientlytoachievetheactualannualyield,but itrarelynearsthegeneticpotential.Reducingtheyieldgap bymoreintensemanagementmaynotbeeconomically

feasibleorenvironmentallyfriendly.Thus,increasing efficiencyofenergy,radiation,nutrient,waterand othernaturalresourcesareobjectives.Theseprocessesare integratedtounderstandandoptimizeplantgrowth,floweringandseeddevelopment.Theintegratedsystemiswhat themanagermustunderstandtoachievethedesiredobjectiveinawaythatissustainablefornowandthefuture.

1

Perspectives,Terminology,and Classification

C.JerryNelson,ProfessorEmeritus, PlantSciences,UniversityofMissouri,Columbia, MO,USA

KennethJ.Moore,DistinguishedProfessor, Agronomy,IowaStateUniversity,Ames, IA,USA

MichaelCollins,ProfessorEmeritus, PlantSciences,UniversityofMissouri,Columbia, MO,USA

DarenD.Redfearn,AssociateProfessor, Agronomy,UniversityofNebraska,Lincoln, NE,USA

Asithasformillennia,theearthischangingphysically, especiallyduringthepastfewdecades,whilehumanpopulationisgrowingveryrapidly.Foragemanagementhas advancedtohelpmeettheexpandingneedsforruminant animalproducts,nitrogenacquisition,fuelresourcesand environmentalstewardship.However,changesinclimate, conflictsandshortagesofwatersupplies,increasedpublic emphasisonecosystemmanagement,andthechallenges ofworldhungerandenergyremaininthenewsalmost daily.Otherconcernsincludefoodsafety,foodqualityand animalwelfare.Eachraisesquestionsabouthowtodeal withhunger,theenvironmentandqualityofhumanlife; especiallyhowmanagementofpastures,foragefieldsand theproductstheysupportcanhelpprovidesolutions.

NeedforConsistentTerminology

Clearcommunicationdependsonterminologythatis commonamongtheindividualsinvolved.Manyterms arecommontoproductionofallcropsandanimals.In thisbook,however,emphasisisonthosetermsuniqueto

foragecrops,pastures,range,andlivestockthatdescribe theirunderlyingscienceandpracticaluse.Termsin boldfacearedefinedinacomprehensiveglossaryinthe appendix.

Whilemanytermshaveahistoryofusage,theycan beconfusingwhenmovedfromonecultureorlocation toanother.Newtermsappearregularlyalongwithnew technologies,andneedtobeclearandusedcorrectly.For example,afewyearsago,adronewouldhavereferredto amalebee,whichitstilldoes,butwiththeadventof precisionagriculture,adroneisalsonowanunmanned aircraftguidedbyremotecontroloronboardcomputers usingglobalpositioningsystems(GPSs).Dronescancarry instrumentsthatmeasureplanthealth,foragequality,forageproductionandmonitoranimalbehaviorinapasture. Manyotherapplicationswillsoonfollow.

Mostdefinitionsarewrittenforthepractitionerand maynotbefullyunderstoodbythegeneralpublicor policymakers.Practitionersaremoreawarethanthe publicorlegislatorsabouttheintrinsicvaluesofforages

Forages:TheScienceofGrasslandAgriculture,VolumeII,SeventhEdition. EditedbyKennethJ.Moore,MichaelCollins,C.JerryNelsonandDarenD.Redfearn. ©2020JohnWiley&SonsLtd.Published2020byJohnWiley&SonsLtd.

andgrasslands.Theyhaveavestedinterestintechnical andeconomicaspectsthathelpthembebettermanagers ormarketers.Somespecificortechnicaltermsforcommunicationamongresearchersareintheglossaryforuse bypractitionersinvolvedwithtechnicalcommunications. Scientists,extensionspecialists,consultants,andjournalistsneedtobeawareofdifferencesinknowledgelevels betweenpractitionersandthepublic,especiallyinurban areas.

TermsinGrasslandManagement

Professionalsinforagesandgrasslandshaveresponsibility todevelopconsistencyofdefinitionssocommunicationis clear.Endophyte-freeorE tallfescue,glandular-haired alfalfa,andno-tillseedingaretermsthatarebecoming common.Conversely,thereisdebateastowhatconstitutesanimalrights,laborlaws,useofwater,andothers, includinghowtomeasurethesefactorsandassignorestimateeconomicvalues.

Whenallowedtodevelopunabated,local,andgeneric termstakeonalocalmeaning.Forexample,thepublic mayobserveapasturethatis“rundown”or“overgrazed.” Thepractitionermightsuggestthepasturewas“grazed heavily,”whereasthescientistmightsayaninappropriate stockingrate, stockingdensity,or grazingpressure, respectively,wasthecause.Eachscientifictermhassome featuresincommonwiththemoregeneraldescriptor, butfocusesonamorespecificfactortoaddclarityusing biologicalreasonsforthepasturecondition.Forexample, overgrazingcouldbeduetopoorplantgrowth,having toomanyanimals,orretainingthemonthepasturetoo long,allwiththeresultofleavingtoolittleresidual forage mass.

“Grazingheavily”suggeststoomanyanimalsforthe forageavailablesuchthattoomuchforagewasremoved. Thescientistwouldusetermssuchas stockingrate (numberofanimalsperunitlandareaforaperiodoftime)and grazingpressure (massofforageavailableperanimalat agiventime)tounderstandthesituationinquantifiable terms.

Sometimesatermusedroutinelyneedstobemodified toleadtochange.“Intensivegrazingmanagement”was commonlyusedfordecadesandgenerallyconnoted theuseofmanagementpracticesinvolving“rotational grazing,”nowcalled rotationalstocking,butitalso impliedthatthe“grazingintensity,”nowcalled stocking rate,wasmanaged.Earlierinterpretationscouldinvolve rotatingperiodsofgrazingandrest,orencouragingfaster biterateorlargerbitesizeofanimals,i.e.“grazingwith intensity.”

Researchonthetechnologiesintroducednewterms thathelpedincreaseproducerinterestinpasturemanagement.Theneedtobebiologicallyaccurate,andconsistent withotherterminologyregardinggrazingmethods,led professionalstoshiftthetermfromintensivegrazing

managementto management-intensivegrazing (Nation 2004).Thisfocusesproperlyonthegrazingmethodthat ismanagedintensivelybasedonknowledgeaboutplants, animals,fencing,watersupplies,andothertechnologies usedasinputs(Gerrish2004).

Inanothercase,professionalsearlyonusedaccumulatedforagefor deferredgrazing;meaningtheforagethat accumulatedduringactivegrowthwasallowedtostand untilneededforgrazing.Yet,practitionersandtechnologytransferspecialistsalsocoinedtheterms“stockpiling” and“grazingon-the-stump,”neitherofwhichwasfunctionallydescriptiveofforageaccumulatedduringactive growth,usuallyduringfall,andsubsequentlygrazedin winterwhengrowthwassloworhadstopped.Evenso,the term stockpiling wasgraduallyaccepted,clearlydefined andisnowwidelyadopted(Figure1.1).

TermsforSoilandItsFunctions

Soilhaslongbeendefinedas“unconsolidatedmineralor organicmaterialontheimmediatesurfaceoftheearth thatservesasthenaturalmediumforthegrowthofland plants.”However,thisdefinitionraisedconcernsamong soilscientiststhatsoilsarenotlimitedtoearth,someparts ofsoilmayberocksorotherconsolidatedmaterial,soils containliquids,gasesandbiologicalorganisms,including plants,andthatsoilsaredynamicduetosoil-formingfactorsthatdifferdependingontheiruseandmanagement (vanEs2017).

UnderleadershipbytheSoilScienceSocietyofAmerica,ideasandconceptswerecoalescedtoanewdefinition: Soil isnow“thelayer(s)ofgenerallyloosemineraland/or organicmaterialthatareaffectedbyphysical,chemical, and/orbiologicalprocessesatorneartheplanetarysurfaceandusuallyholdsliquids,gasesandbiotaandsupport plants”(vanEs2017).Thenewdefinitionclearlyplaces moreemphasisonthephysicalmakeupofthesoiland broadensthedefinitionandusesbeyondagriculture,i.e. morethanjustsupportingplants.

SoilQuality

Anumberofyearsago,thetermsoilqualitywasintroducedandconsideredas“thecapacity(ofsoil)tofunction” (Karlenetal.2003).Soilqualitydependsonphysical, chemical,andbiologicalfeaturesofupperlayers,and howtheyinteracttoprovideagivenfunction,beitfor roadconstruction,cropproductionorahomelawn.A changeinonefeatureresultsinadifferentsoil.Scientists aredevelopingmethodstoassesstheindicators,andthen usemathematicalequationstocombineseveralphysical, chemical,andbiologicalfeatures,includingorganic matterthatchangeswithhumanactivity,intoanumeric index(Friedmanetal.2001).Thedesirednumericindex basedonphysical,chemical,andbiologicalproperties wouldvarydependingonthepurpose,e.g.agricultureor civilengineering.

FIG.1.1. BeefcattleinSaskatchewanextendingthegrazingseasonbyusingaccumulatedforage.

Source: PhotocourtesyofVernBaron.

Understandinghowthecomponentsofthesoilquality indexinteract,whilebeinganoblegoal,hasbeendifficult tomeasureandinterpretoverarangeofsoiltypesand topographies(Laishrametal.2012).Thisledtointerest insoilhealth,asimplerconceptforevaluating“soilvalue” thatisrelatedmoredirectlytocontentoforganicmatter (DoranandZeiss2000).Thisseemedmorepractical foragriculturaluses,especiallyintheshortterm,since organicmatterisresponsivetomanagementandaffects thestructureofthesoilanditscapacityforholdingwater andnutrients.

SoilHealth

Soilhealth isthecontinuedcapacityofsoiltofunction asavitallivingecosystemthatsustainsplants,animals, andhumans.Thetermmaybemoreusefulthansoilqualitytodescribethe“healthstate”ofasoilintermsofits productivityandrolesinenvironmentalconservationand themanyecosystemservicesofpasturesandforagessince itisbasedmainlyonorganicmattercontentthatisrelativelyeasytomeasureandquantify.Manypractitioners andtheUSDANaturalResourcesConservationServices

haveadoptedthetermsoilhealthusingorganicmatteras themaincomponentforevaluatingsoilconditionsassociatedwithcropandforagemanagement.Soilhealthand itsemphasisonorganicmatteristhetermchampionedby organicagriculturalists.

SustainabilityofGrasslandAgriculture

Sustainabilityofagriculturehasbeenacriticalissuefor farmersandranchersforgenerations,butinthe1960’s publicconcerngrewabouttheincreasedemphasisonprimaryproductionoffoodandfiberbasedonuseofchemicalfertilizersandpesticides.Agriculturewasperceivedas miningnaturalresourcesforeconomicbenefitwithlittle concernforshort-andlong-termsustainabilitybasedon healthandwell-beingofconsumersandtheenvironment. Thepublicraisedrealconcernsaboutgovernmentregulationsandmanagementpracticesforuseofchemicalson farms.

TheDelaneyamendmentin1958prohibitedany compoundinfeedsorfoodsthatcausedcancerinanimals orhumans.Tohelpmeettheseconcerns,thegovernment developedstricterregulationsonuseofchemicalfertilizers

andespeciallypesticidesbasedonbetterdiagnosticprocedures.Thiswascoupledwithincreasedpublicinterest inorganicagriculturethatprohibitsuseofchemical inputs.Toaddressthegrowingconcerns,theconcept ofsustainableagricultureemergedand,earlyon,was somewhatlinkedtotheuseoforganicpractices.

RolesofthePublic

Thepublicwasconcernedaboutsustainabilityeven thoughitwasclearorganicagriculturealonecouldnot meetthetotalfoodneedsthen(Peseketal.1993)orlater (ReganoldandWachter2016).Soonamoreholisticperspectiveofsustainableagricultureemergedthatinvolved morethanfoodproductionandwasdefinedbasedon threemajorcomponents:(i)theeconomicreturntothe producer,(ii)theconservationoftheenvironment,and (iii)useofpracticesthatareacceptedsocially(American SocietyofAgronomy1989).

Federalandstategovernmentsbegancost-shareprogramstoencourageandrewardproducerswhoadopted managementpracticestoreducesoilerosion,maintain waterquality,increaseplantdiversity,enhancewildlife andreducenegativeeffectsofchemicalnutrientsandpesticides.Industryalsoacceptedthechallengesbyworking onthebroaderissuesbeforesubmittingchemicalsfor registration.

Today,thereisgrowingconcernaboutsocialaspects likeanimalrights,workersafety,foodsafetyandlabelingofcontentsinfoodascomponentsofsustainability. Inmanycasestoday,theconsumercangetsomereassurancebypurchasingfooddirectlyfromFarmer’sMarkets ortrackproductsbacktothefarmorranchfromwhichit wasproduced.

ConsiderationofEcosystemServices

Afteradetailedinternationalanalysis(Millennium EcosystemAssessment2005),sustainabilityofagriculture todayalsoincludesprovidingawiderangeof ecosystem services,amoreinclusiveandmorecomprehensivesetof ecosystemcomponentsandinteractionsaffectinghuman well-being.Thisfour-partframework,ledmainlyby ecologistsandsocialscientists,consistedoffouroutputs orservicesfromtheland(Figure1.2).

Thedesiredoutputs,allexpectedfromagriculture, include(i)Supportingserviceslikeprimaryproduction, nutrientcyclingandsoilformation;(ii)Provisioning serviceslikefood,freshwater,wood,andfuel;(iii)Regulatingserviceslikeinfluencesonclimate,quantityand qualityofwater,anddiseasesofplantsandanimals;and (iv)Culturalserviceslikespiritualissues,education,and esthetics.Currently,amajorgoalforscientistsistolearn thebreadthanddeterminevaluesofindividualecosystem servicesandtheirinterrelationships.

Themillenniumreportonsustainableagricultureis graduallybeingacceptedinternationallyasagoal,while

moresub-componentsareaddedonaregularbasis. Costanzaetal.(2017)reportedontheexplosionof researchbyecologistsandeconomistswantingtoassign valuestoecosystems,encouragepoliciesanddocument applicationsoftheideas.Sustainabilitynowextends beyondthefarmgatetotheentirefoodchainand includesamyriadofenvironmental,social,andcultural issuesrarelyconsideredafewdecadesago.Agricultural scientistsneedtocontinuetobeinvolvedinallaspects.

Unfortunately,agriculturalsciencehasoftennotkept uptoprovideascientificbasisforleadershiptomakegood policydecisions.Thepublicisnowthemajorplayer,often withoutscientificevidence,indecisionsandregulations fortheentirefoodsystemandthepreservationofnatural resources.

Assessingandunderstandingthecomplexityinvolved withagriculturalsustainabilitywilllikelyrequiremathematicalmodelingandtransdisciplinaryapproaches inresearch.Forageandpasturemanagementandanimalwelfareissuesneedscience-basedcooperationwith socialscientistsandpractitionerstounderstandrelationships,provideeducationandsatisfypublicdemandsfor sustainability.

Industrializationofagriculturevianewtechnologies fromboththepublicandprivatesectorshasraised concernsaboutethicalandeconomicmotivationamong theplayers.Thequestionarises;arecommercialmotives parallelwiththoseofthepublic,andbasedonscience? Scientistsareintheearlystagesofestablishinganindex thatincludesmeasurablevariablesassociatedwiththe MillenniumAssessmenttoachievesustainabilityinways thataresociallyacceptable.Asincomesincreaseindevelopedcountries,demandsforfreshandsafefoodswith goodtastewillcontinuetorise.Manywillbelieve,with littleornoscientificevidence,thatorganicallyproduced foodsaresafer,healthier,andtastebetter.Thebalance betweenorganicandotherproductionsystemswillevolve (Tillmanetal.2002).

TheRoleofOrganicAgriculture

Organicfoodsandbeveragesareasmall,butrapidly growingmarketsegmentintheglobalfoodindustry includingmeatandmilkproducts,primarilydueto healthandnutritionconcerns.Arecentstudyanalyzed 40yearsofsciencecomparingorganicandconventional agricultureacrossthefourgoalsofsustainability,productivity,environmentalimpact,economicviability,and socialwell-being(CrowderandReganold2015).Insummary,organicsystemsproducedloweryieldscompared withconventionalagriculture,yetitwasmoreprofitable becauseconsumerspay12–50%morefortheproducts.

Overall,organicfarmstendtostoremoresoilcarbon, havebettersoilquality,andreducedsoilerosion.Initialevidenceindicatesthatorganicagriculturalsystems delivergreaterecosystemservicesandsocialbenefits.

FIG.1.2. Left,ecosystemservicesaredividedintofourboxesrelatedtomajorserviceswithineach. Right,listofcomponentsofwell-being.Linedarknessindicatesthepotentialeffectsofsocioeconomic factorswhereaslinethicknessshowsintensityofeffectsofecosystemservicesonhumanwell-being. Source: FromMillenniumEcosystemAssessment(2005),presentation;CreditedtoMillenniumEcosystemAssessment(2005).

Althoughorganicagriculturehasanuntappedroleto playwhenitcomestotheestablishmentofsustainable farmingsystems,nosingleapproachwillsafelyfeedthe planet.Rather,ablendoforganicandotherinnovative farmingsystemsisneeded.Significantbarriersexistto understandingandadoptingthesesystems,andadiversity ofpolicieswillberequiredtofacilitatetheirdevelopment andimplementation.

Amajorsocialissueisthatorganicagricultureisconceptuallyassociatedwithsmallfarmswithamixofcrops andlivestockthatareownedoroperatedbyafamily.Also, organicproductsareoftenmarketedinnearbyfarmer’s marketswherefreshnessandrelationshipswiththeproducerarevalued.Duetoincreasingpublicdemandfor organicproducts,theyareofferedinmostsupermarketsat pricesfrom15%to50%ormorethanproductsproduced traditionally.

Incontrastwithorganicplantagriculture,morethan 90%oflivestockproductsareproducedonlargefarms

usingtraditionalpracticesforplantandanimalmanagementthattendtobefocusedononlyoneortwocommodities.Mostareownedandmanagedbyafamily,but haveseveralemployeeswhodomuchofthework.Due tosize,theseoperationsaremoreeconomicthansmaller farmsbecauseinputsareoftenpurchaseddirectlyfrom suppliersandproductsaremarketedthroughpre-arranged contractstoobtainahigherprice.Thisleadstogreaternet incomeforthefamilyorowner.

Manyconsumershavenegativeperceptionsofproductionagriculturethatishighlymechanizedwithlargefields thatdeterwildlifeorhasahighdensityoffarmanimals. Theycriticizeuseofgeneticallymodifiedplantsoffered byprivateindustry,safety-approvedpesticides,economic ratesofchemicalfertilizers,andconfinementhousing foranimals,evenifallpracticescomplywithfederal regulations.Operatorsofverylargefarmsareaccused ofexploitinggovernmentassistanceprogramsanddisregardinganimalrights,workerwelfareandenvironmental

regulations.Consumerdemandinthelongtermwill determinetheproportionoffoodsproducedbyorganic ortraditionalmeansoneithersmallorlargefarms.

ClassificationSystemsBasedonCropUse

Somesystemsofplantclassificationaroseoutofconveniencewhileothersaroseoutofnecessity,suchasthe binomialsystemofplantnomenclaturebasedonmorphologicfeatures.Newclassificationcategoriescontinue toemergeastechnologiesanduseschange.Forexample, a medicinalcrop isgrownforitsnaturalproductsfrom theleaves,flowersorrootsthatareusedformedical purposes.A pharmaceuticalcrop, sometimesgenetically engineered,isgrownprimarilyasabiologicsynthesizerof aspecificcompoundusedformedicalpurposes.

TermsforAgronomicUses

Agronomy,derivedfromtheGreektermfor“field,”deals withfieldcropsincludingwheat,corn,soybean,cotton, andforages.Thesearegrownonalargescaleusingrelativelyextensivemanagementcomparedwithhorticultural crops. Forage includesediblepartsofplants,otherthan separatedgrain,thatcanprovidefeedforanimals,orthat canbeharvestedfor“feeding.”Thus,itincludesleaves, twigs,stems,roots,nuts,andotherpartsofawiderange ofplantspecies.

Primaryusesofforagesassociatedwithfeedforanimals are pasture, hay, silage,and soilage.Pastureisa grazing managementunit thatisenclosedandseparatedfrom otherareasbyfencingorotherbarriersandismanaged toproduceforagethatisharvestedprimarilybygrazing. Rangeislandsupportingnativevegetationthatisgrazed orhasthepotentialtobegrazedand,incontrasttopasture,isusuallymanagedextensivelyasanaturalecosystem. Inadditiontograsses,legumes,andother forbs (Smith andCollins2003),rangeincludesshrubsandtreesthat provide browse foranimals.

Hayisforagepreservedbyfielddryingtomoisturelevelslowenoughtopreventmicrobialactivitythatleadsto spoilage.Incontrast,silageisforagepreservedinasucculentconditionatlowpHduetomicrobialproductionof organicacidsbyanaerobicfermentationof sugars inthe forage.Soilageorgreenchopisforagethatiscutandfed freshwithinafewhours.Browseisleafandtwiggrowthof shrubs,woodyvines,trees,cacti,andothervegetationthat isavailablefordirectanimalconsumptionby“browsing.” Acatchcropisaforagecrop,usuallyanannuallikesudangrassthatisusedshort-terminarotationwithoneormore rowcrops.Catchcropsareusedwhenseverewinterinjury orothersituationarisesandmoreforageisneededinthe short-term.

Insomecases,foragespeciesaregrownforprimarypurposesotherthananimalfeed.A greenmanurecrop is allowedtoproducevegetationtobetilledunderforsoil improvement;a grassedwaterway isplantedinsurface

drainageareasoflargefieldstoacceptsurfacewaterand channelitoffthefieldtoreduceerosionandgullyformation.A smothercrop isastronglycompetitivecropthat isgrowninmonoculturetocontrolweedsuntilitisharvestedorgrazed.A companioncrop (thepreferredterm over nursecrop),suchasoatorspringwheat,canbesown atareducedratealongwithaforagecropthatemergesand developsslowly.Thecompanioncropestablishesquickly toreduceerosionandcompetewithweeds.Inallthese cases,theforageorgraincanbeharvestedifremovaldoes notinterferewiththeprimaryobjective.

Covercrops areusedtostabilizethetopsoilandreduce waterrunoffanderosionbetweensuccessiveannualcrops, oftenoverwinter(Finneyetal.2017).Usually,awinter grain,winterlegumeorrootcroplikeradishorturnipis plantedinautumnafterthepreviouscropwasharvested. Growthofrootshelpholdthesoilparticlestogetherwhile thetopsinterceptrainfallandreduceimpactofwater dropletsthatcandislodgesoilparticles.Theforagecan beharvestedinspringorkilledtoleavemulchfordirect plantingofthenextcrop.

DetailedstudiesinPennsylvaniaindicatedpositive andnegativeeffectsfromdifferentcovercrops.Legumes areusuallypreferredbecausetheyfixsomeN,whereasN appliedtograssremainssequesteredinthekilledtissue duetoslowmineralization,canleadtolowNsupplyand lowyieldsinthesubsequentcrop(Whiteetal.2017).But yieldsandotherecosystemserviceslikeweedandinsect controlwerebetterwithnon-legumespecies,sotradeoffs needtobeconsidered.Also,changesinplantingdates andseedingratesalteredtheecosystemvaluesofcover crops(Murrelletal.2017).

TermsforEconomicLandUses

Croplandforage iscultivatedinsomeway,usuallyas partofarotationwithagrain,fiber,oroilseedcropthat includesforagesandshort-termpastures.Forageshelp controlerosion,increasesoilorganicmatter,improve aeration,andlegumesleaveresidualNinthesoilfor subsequentcrops.Croplandforages,includingcornstalks orothercropresiduesthatarepartofacroprotation,can beharvestedforhayorsilage,orcanbegrazedaspastures. Theseareasalsoserveassitesforapplicationofmanures.

Grazingland includesbothpasturelandandrangeland,theformermorecommoninthehumidareasof NorthAmericaeastofthe98∘ meridian,usingintroducedforagespeciesinsystemsthataremoreintensively managed. Rangeland consistslargelyofnativespeciesin thesemi-aridwesternpartsofNorthAmericathatare managedmoreextensively.Nativegrasslandspeciesare moredroughttolerant,usuallylowerinherbageyield, andmoresensitivetograzingmanagementthanare the introducedspecies thatpredominateintheEast. Availabilityofwaterandcompetitionwithothercrop speciesforlanduseintheEastoftenrelegateforagesand

pasturestolandclassesthatarelessproductiveortoo erosiveforcropproduction.

Forestland consistsofsomewhatopen,tree-covered areasthatsupportforageandgrasslandspeciesthatcan begrazed(Garrettetal.2000)orbrowsed(Figure1.3). Grazingofferssomeanimalproductionandhelpscontrol understoryvegetation.ThesesystemsintheWestcan expanduseofforestland,orintheEastcanaddafew yearsofcroporforageusebeforethetreecanopycloses inaplannedapproachtoforestmanagement.These systems,ofparticularimportanceinthepineforestsof thesouthernandsoutheasternUS,arerelativelycomplex todesignandmanage,butcanbeveryproductive(Child andPearson1995).

Agroforestry isadesignedmanagementsystemin whichtreesarepurposelyspacedtoallowplantingof cropsorforagesamongthemorinalleysbetweentree rows.Thecombinedobjectivesareshort-termanimalor cropproductionfromthealleysforafewyearsfollowed byintermediate-termincomefromnutsforfoodorneedleproductionformulchuntilthetimberisharvested. Theseareoftenusedwithproductionofhigh-valuetrees suchaswalnutorpecanthatproducenutcropseach year.Thisagronomicuseforthefirstyears,provides erosioncontrol,Nfortheecosystemiflegumesareused, increasedbiodiversityoftheareaandhabitatoptionsfor wildlife.

Silvopasture systemsareagroforestrysystemsusing pastureplantsthatoccupyopenareasoralleysbetween treerowsthatcanbegrazed(ClasonandSharrow2000). Theyoungtreesneedprotectionfromdamage,becared

forandprunedregularly(Figure1.4).Theultimategoal istoprovideincomefromlivestockproductsintheshort termwhilethecanopygraduallyclosesandprovides toomuchshade.Thelonger-termgoalofthesystemis toproducenutsorotherproductsuntiltheharvestof well-shapedtreesofhighvalue.

TermsforEcologicalLandUses

Foragesandgrasslandsplaymajorrolesinenvironmental stabilitybyreducingerosion(seeChapter12andSharp etal.1995),improvingwaterquality(Chapter12), increasingbiodiversity(Chapter13),andprovidingfood andhabitatforwildlife(Clubine1995;Sollenberger etal.2012).TheConservationReserveProgram,afederalprogramtopayUSlandownerstoremovehighly erosivelandsfromcropproduction,isbasedonthese principles.The10-yearcontractrequireslandmanagers toplantadaptedperennialforagespeciestomaintain year-roundgroundcover.Theresultisreducedwater runoff,enhancedwaterqualityandimprovedwildlife habitatontheconservedland.Inaddition,theprogram helpsreduceoverproductionofcropsandtheneedfor subsidypaymentsbythegovernment.

Grassedwaterways providedrainagechannelsfor cropfieldswhereasplantingforagesin riparianbuffers protectingstreamshelpscontrolsoilerosionandcapture nutrientsandothermaterialscarriedinrunoffwater (Chapter12).Desiredwidthsofwaterwaysandriparian stripsdependonscientificestimatesofexpectedratesand volumesofrunoff.Waterwaysandriparianareascanbe harvestedforhayorsilageatappropriatetimesduringthe

FIG.1.3. Aforestlandpasturesysteminwhichtreesshadethepastureandcattle.Thelargetrees resistanimaldamageandarespacedtoproducequalitytimber. Source: PhotocourtesyofNational AgroforestryCenter.

FIG.1.4. Asilvopastoralsystemwithwinterryeplantedamongtreesandinthealley.Othercrops canbegrowninthealleysuntilthetreesgetlargerandalleysgraduallynarrow. Source: Photocourtesy ofRobKallenbach. growingseason,ifadequatestubbleisleftforsubsequent regrowthtostillprovidetheneededprotection.

RelationshiptoPrecipitation

Theecologicbasisforlandtypesdependsonclimatic factorssuchasprecipitationandsoilmoisture. Desert isobviouslyanaridlandclassification.Movingtoward areasofhigherandhigherprecipitation,ortoareaswhere evapotranspiration decreases,thenaturalvegetation progressivelyphasesto shrubland, steppe, and prairie (Chapter8).Desertplantsoftenhavecrassulaceanacid metabolisminwhichstomataopenforCO2 uptake onlyatnighttoconservewaterandtoleratedrought(see Chapter4).Inaddition,severalshrubsavoidherbivory duetospinesortastefactors(Chapter46)tosurvive anddominateinadryarea.Steppesusuallyoccupydrier areasthanprairieandconsistmainlyofdeep-rootedshort grasses.EasternportionsoftheprairiesinNorthAmerica consistnaturallyoftallgrasses. Range ismoreencompassingandincludesareassuchasdesert,shrubland, steppes,andprairie.

Inhighprecipitationareas,unlessburnedormanaged correctly,thenaturalgrasslandvegetationwillgradually beovercomebywoodedvegetationorforest. Marshland and wetlands areareasofhighprecipitationorpoorsoil drainagesuchthatahigh-watertableismaintainedfor

muchoforallyear.Theseareascanbegrazedwhensufficientlydry,butserveprimarilytoreduceflooding,providewildlifehabitatandmaintainbiodiversity. Meadows aregrasslandsites,oftenwithnativeornaturalizedspecies thatexistaslong-termstands,butproductivityisaffected stronglybythelandscapetopographyandwater-holding capacity.Assuch,theyareoftennaturallysub-irrigatedin theWestordependonnaturalrainfallintheEast.They areusuallygrazedorharvestedforhayduringdryperiods. Oftenanadjectivesuchashay,mountain,native,orwetis usedtohelpdescribethemeadow,i.e.mountainmeadow.

RelationshiptoTemperature

InNorthAmerica,highairtemperaturesinJulyandlow airtemperaturesinJanuaryareprimaryfactorsaffecting adaptationofgrasslandspecies(Chapter8). Tundra is treelessgrasslandthatoccupieslargeareasofarcticregions ofNorthAmerica,Asia,andEurope.Inwarmerareas, plantswithgoodwinterhardinessandactivegrowth atlowtemperatures,i.e.cool-seasonspecieswithC3 photosynthesis(Chapter4),dominateeasterntemperate grasslands.Duetohightemperaturesanddryconditions, however,warm-seasonspecieswithC4 photosynthesis andgoodwinterhardinessgrowactivelyinsummerand dominatemanytemperateregionsoftheNorthAmerican prairie.

Furthersouth,the transitionzone consistsofsome areasofcool-seasonspecieswithC3 photosynthesis,and otherswithwarm-seasonspecieswithC4 photosynthesis. Highsummertemperaturesrestrictmanycool-season grassesinthisareaespeciallywhengrazedtoashortstubbleheightwhensoiltemperaturesarehigh.Conversely, severalnativewarm-seasongrassesarehighlyproductive andsomesubtropicalgrassescansurvivethemilder winters.

Dependingonthemildnessofwintertemperatures, subtropicalperennialgrassesandsome herbaceous legumesoccupythegulfregion.Farthersouth,most foragesaretropicalspeciesthattolerateheat,butarevery sensitivetocold(Chapter18).Woodyplants,especially treelegumessuchasleucaena,canbeavaluableforage componentinsubtropicalandtropicalareas(Chapter15). Savannas describegrasslandswithscatteredtrees,often legumetreesinthetropicsandsubtropics,orhardwood treesintemperateareas.

TermsDescribingVegetationTypes

Forages,rangelands,andpasturesconsistofvegetation thatcoexistsindifferentstagesorconditions.Plantfunctionaltypesinadiversemixtureconsistofgrasses,forbs, brushorshrubs,andtrees.Eachprovidesafoodsource forharvestordirectgrazingbyanimals.Inthebroad sense, forbs are herbaceous (non-woody)broad-leafed plantsthatincludethelegumes(SmithandCollins2003). However,whendescribingforages,legumesareusually consideredseparatelyfromforbsduetotheirhighereconomicvalue.Thus,ingeneral,forbreferstonon-legume, herbaceous,broad-leafedplantssuchasdandelionand several Brassicas includingrape,turnip,andkale.

Forbsincludesomenaturallyoccurringpoisonous plantsandotherscommonlyconsideredasweeds,but foragequalityofseveral“weeds”isasgoodand,insome cases,evenbetterthanseededspecies(Martenetal.1987). Someforbssuchasdandelionandgoldenrodareinvasive andneedtobekeptincheckwithgoodmanagement (Chapter28).Manyforbsknowntobeweedswithgood foragequalityarestrongcompetitorswithcropplants. Therefore,theyareconsideredundesirableinpastures andforagefieldsnearbyorinarotationwithcropspecies. Thepositiverolesandpotentialnegativeconsequencesof theseforbsneedfurtherevaluation.

Foragecanbeseededandharvestedasa monoculture, i.e.asinglespecies,orasamixtureoftwoormorespecies. Herbage, theabovegroundmaterialthatconsistsof leavesandstems,usuallyrefersto foragemass harvested mechanically,whereas forageavailable referstothe massthatcanbegrazedtoadefinedheight. Aftermath describestheregrowthafterharvest,whichcanbehigh quality,ortheresidueleftinthefieldafterseedharvest thatisusuallylowquality.

Stockpiled forageresultsfromaspecialmanagement strategytograzetheaftermathordeferredgrowthof cool-seasonorwarm-seasongrassesduringapartofthe yearwhenplantsarenolongergrowingrapidly.Stockpilingisusedregularlytoaccumulatevegetative(leafy) growthofcool-seasonspeciesduringlatesummerand falltoextendthegrazingseasonintothewinter-dormant period(seeFigure1.1).Thiscanbeacost-savingalternativetoreducefeedingconservedforageashayorsilagethat requiresharvestandsomeformofstorage(Chapter20).

TermsDescribingLifeCyclesandStandPersistence

Someforageandgrasslandplantsare annuals that completetheirlifecycleinoneyear. Summerannuals germinateinspring,growactively,produceseed,and thendie.Somedieasadirectresultof flowering andseed productionthattriggersacoordinatedandprogrammed death.Thisinvolvesreallocationoforganicandmineral resourcesfromthestem,root,andleavestotheseed. Othersummerannuals,suchascrabgrassandannual lespedezas,continuetogrow,inanindeterminatemanner afterfloweringandproducingseed,untilkilledbycold temperature.

Sudangrassisasummerannualthatflowersinsummerbutdiffersfromcorninthat,likecrabgrass,ittillers activelyandregrowsafterfloweringorcutting.Sudangrass eventuallydiesbecausethissubtropicalgrassissensitiveto frostanddoesnotdevelopwinterhardiness.Standpersistenceofsummerannualslikekoreanlespedezaorcrabgrassdependontheirabilitytoproduceseedthatmust overwintertogerminatethefollowingspring(Beuselinck etal.1994).

Winterannualslikecrimsonclovergerminateinfall, growvegetativelyoverwinter,andthendieafterflowering thefollowingspring.Theygenerallyhaveprogrammed senescence processesthatbeginshortlyafterflowering andseedproduction.Theydependonseedsurvivalover summertogerminatethefollowingfalltoprovidestand persistence.

Someforbs,includingsweetclover,aretrue biennials Theygerminateinspringandremainvegetativebyproducingonlyleavesandstemsandformalargetaproot duringthefirstyear.Afteracold-inductionperiodduringwinter,theyflowerandproduceseedinspring.They havelittle,ifany,storageoforganicresourcesintheroot orcrowninyeartwo,producelittleornoregrowthand donotsurvivethesecondwinter.Sincetheplantssurvive onlytwoseasons,long-termstandpersistencedependson seedproductionandseedlingdevelopment.Sweetclover hasadaptedbyhavingahighpercentageof hardseed, someofwhichdoesnotgerminateforseveralyears.No knowngrassisatruebiennial.

Standpersistence,thelongevityofaplanting,can dependoninnatelongevityoftheseedlingsthatbecome establishedplantsthatsurvive(i.e.plantpersistence)or

theabilityofshort-livedplantstospreadvegetativelyor byseed(Beuselincketal.1994).Plantslikealfalfaand sericealespedezaarelong-lived crown formerswitha seedlingrootthatsurvivesforseveralyearsandmaintains acrownofbudsatthesoilsurfaceforoverwintering andregrowth(Chapter3).Birdsfoottrefoil,alsoacrown former,isintermediateinthattheseedlingssurviveforup tofouryearsincoolenvironmentsbutlessthantwoyears inhotenvironments,sostandpersistencedependson bothplantsurvivalandreseeding.

Incontrast, clone formerslikewhitecloversurviveand spreadby stolons or rhizomes thatbothformnewroots atnodesandproducenewshootsfromaxillarybudsat rootednodes.Thus,whentheseedlingrootdies,thenewly rootedstolonsorrhizomesoftheclonalplantcontinueto growandproducevegetationtoperennate.

TermsInvolvingBiodiversity

Mixedswardsallowspecieswithdifferentgrowthpatterns andmaturitytominimizediseaseandpestproblemsof monoculturesandhelpbalanceproductionratesthroughouttheyear.Severalstudieshavedemonstratedthepositivevalueofspeciesdiversityonproduction,especially innaturalecosystems(Loreauetal.2002;Figure1.5). Biodiversity referstothenumberofspeciesorfunctional groupsinahabitat,whichhasaneffectonseveralkey ecologicprocessesincludingbiomassproductivity,ratesof mineralizationofsoilnutrients,andstabilityorlongevity ofthesystem.

Biodiversityofseveralspeciesisusuallycharacterized asafunctionofspeciesrichness,i.e.thetotalnumber ofspeciespresent,andtheproportionalabundanceof eachspecieswithinthecommunity.Evennessrefers tothedistributionofspecies;highevennessindicates theproportionsofspeciesaresimilarorhomogeneous withinthecanopy.Inmanydiverseplantcommunities,

FIG.1.6. Productivityofindividualspeciesin amixed-speciespastureorgrassland.Most productioncomesfromafewspeciesthat dominatethestand.Thoughnothighly productiveduetolowdensityorlowyieldper plant,theremainingspeciescontributeto stabilityandresilienceoftheecosystem. Source: CompiledfromseveraldatasetsfromtheGreat Plains. arelativelyfewspeciespredominate(Figure1.6).These communitieshaveaheterogeneousorunevendistribution ofspeciesand,therefore,lowevenness.Theinverseof evennessisdominance,soplantcommunitieswithlow evennesshavehighdominanceandviceversa(Peet1974).

Plantscanbeclassifiedintofunctionaltypes,based eitherontheirresponsesoreffects.Response-functional typesconsistofplantspeciesthatrespondinasimilar mannertoabioticandbioticconditions.Ingeneral, sincethespecieshavesimilarfunctionaltraits,theyare theoreticallyinterchangeableintheplantcommunity.For example,theymayhavesimilarreproductivestrategies, growthhabits,orcarbonmetabolismthatenablethemto persistinthepopulationinresponsetoherbageremoval bycuttingorgrazing.

FIG.1.5. Theeffectofspeciesrichness (numberofspeciespresent)onproductivity ofanaturalgrassland. Source: Compiled fromseveraldatasetsfromtheGreatPlains.

Effect-functionaltypesareplantspeciesthataltertheir processessuchastheirproductivityornutrientcyclingin asimilarway.Typesthatalterproductivityinasimilarway couldincludeC3 grasses,C4 grasses,legumes,non-legume forbs,andwoodyspecies.Forexample,inNorthAmericangrasslandsandshrublands,temperatureisusuallythe mainfactoraffectingabundanceofC3 versusC4 grasses (ParueloandLauenroth2005).C4 grassesarefavoredby warmerclimatesinthesoutheasternUS.Abundanceof forbsislessaffectedbygeographicandclimaticvariables, butthespeciesofforbinthepopulationcanchange.

Highproductivityofnaturalgrasslandecosystemsin westernareaswheresoilfertilityand/orsoilmoistureare limiting(Huston1994)isoftenassociatedwithincreased

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