5/10/2017 HydrofarmAReviewofFactorsAffectingPlantGrowth

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Lighting A Review of Factors Affecting Plant Growth

Hydroponics MarianneAmes,GraduateFellow
WayneS.Johnson,AssistantProfessor
ClimateControl UniversityofNevada,Reno

GardenAccessories Growthisdefinedasanirreversiblechangeinthesizeofacell,organorwholeorganism.Itmayalsobetheincreaseincell
numberwithoutchangesinvolumeorweight.Commonly,growthistheincreaseintheamountoflivingmaterial(protoplasm)
GrowingMedia
whichleadstoanincreaseincellsizeandultimatelycelldivision.Theincreaseinprotoplasmisbroughtaboutaswater,carbon
dioxideandinorganicsaltsaretransformedintolivingmaterial.Growthoccursonlyinlivingcellsbymetabolicprocessesinvolved
PlantCare
inthesynthesisofproteins,nucleicacids,lipids,andcarbohydratesattheexpenseofmetabolicenergyprovidedby
photosynthesisandrespiration(7).
SeedStarting

Nutrients&Additives Differentiationistheprocessesinvolvedintheestablishmentoflocalizeddifferencesinbiochemicalandmetabolicactivityandin
structuralorganizationthatresultinnewpatternsofgrowth(7).Differentiationofindividualcellsinvolvesthesystematicturning
FoodStorage onandoffofgeneticcontrolmechanisms,withmitoticprocessesincelldivisioninsuringgeneticcontinuityofallcells(7).

Books&Videos Wholeplantdevelopmentistheorderlyandprogressivechangefromseedgerminationthroughjuvenility,maturity,floweringand
fruiting.Environmentalfactorsmayinfluencedevelopmentaltimesorblockparticularstagesaltogether(7).
Aquaponics
Thetermsgrowth,differentiationanddevelopmentencompasstheeventsrelatedtotheprogressiveunfoldingoftheplantsgenetic
informationinrelationtoenvironmentalcues.Theplantreceivesvariousinternalandexternalstimulithatinteractwiththegenetic
informationwhichmaythenchangemetabolicactivityandinfluencestructuralorganization(12).
By Brand
Growthinplantsisrestrictedtocertainembryonicregions,variouslycalledmeristems,budsandcambium.Theselocalized
selectone
embryonicregionsofhigherplantsdifferfromanimalswheregrowthtypicallyoccursthroughouttheorganism.Thestemsand
rootshaveapicalmeristemsresponsibleforextensiongrowthwhichusuallyremainpermanentlyembryonicandcapableofgrowth
forlongperiods.Growthingirthoccurswithcelldivisionandenlargementincambialtissueofstems(16).
New Arrivals
Thegrowthanddevelopmentalpatternsofplantsarecommonlyusedtoclassifyplantsintogroups.Annualplantscompletetheir
entirelifecycle,fromseedtoseed,inasinglegrowingseason,whereasbiennialplantsrequiretwogrowingseasons.Perennials
growyearafteryearoftentakingyearstomature.Inherbaceousperennialstherootsandshootscanremainaliveindefinatelybut
theshootsystemmaybekilledbyfrost.Eachspringshootgrowthresumesfromadventitiousbudsatthecrownoftheplant.In
woodyperennials,boththeshootsandtherootsremainaliveindefinitely(7).

Indeterminateplantsarethosewhosemainaxesremainvegetativeandinwhichflowersforminaxillarybuds.Theseplantshave
shootswhichcontinuetogrowinfavorableconditionsandincludesuchviningplantsascucumbers,peasandgrapes.A
determinateplant'smainandsecondaryaxesterminateinaflowerbudandconsequentlyshootelongationstopsasinsweetcorn,
bushtomatoes,peppers,bushbeans,etc.(4).

MeasurementofGrowth

Plantgrowthisoftenmeasuredasachangeinarea,length,volume,height,wetordryweight.Thesemethodsmaynotalwaysbe
asatisfactorymeasureofgrowthataparticularstageofplantdevelopment,i.e.,agerminatingseedmayshowanoveralllossin
dryweightduetotheutilizationoffoodreservesduringrespiration,althoughtheseedisdefinitelygrowingasevidencedbyits
emergingrootsandshoots.

Therelativegrowthrate(RGR)whichisthesizeincreaseperunitintervaloftimehastwocomponents:thenetassimilationrate
(NAR)andtheleafarearatio(LAR).TheNARistherateofincreaseofdryweightperunittimeperunitofleafsurfacewhichisa
measureoftheamountofphotosyntheticproductgoingintoplantmaterial.TheLARistheratioofleafareatodryweightwhichis
themeasureoftheproportionoftheplantthatisengagedinphotosynthesis(12).Combinedtheygivearelativedescriptionof
growthovertimebaseduponplantcharacteristics.

VegetativeGrowth

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Germinationincludesallthestepsfromtheseedimbibingwateruntiltheseedlingisselfsustaining.Withintheseed,reserve
substancesareenzymaticallyconvertedintomaterialsusedinsynthesisoroxidizedthroughrespirationtoreleaseenergy.The
seedrequireswater,air(oxygen),andthepropertemperaturerangesuchthatbiochemicalprocessescanoperate(7).

Aseedisconsideredgerminatedwhenithasproducedaplantthatispotentiallycapableofcontinuousgrowth.Fromthebeginning
ofthisstage,untilinitiationofthefirstflowerprimordium,theplantisinthevegetativestageofgrowth.Whenaplantcannotbe
madetofloweritissaidtobejuvenile(7).

Thejuvenilegrowthphaseischaracterizedbythemostrapidrateofgrowththeplantwillundergo.Aswell,thejuvenileplantmay
exhibitdifferentmorphologicalorphysiologicalfeaturesthanamatureplantofthesamespecies.Acommonfeatureofmany
juvenileplantsistheabilitytoinitiateadventitiousrootsreadily,anabilitywhichisoftendecreasedorlostinmatureplants.The
juvenilephasevariesfromonetotwomonthsforannuals,tomanyyearsforwoodyperennials(7).Theabilitytoinfluencethe
lengthoftimeaplantisinthejuvenilephaseisimportantinsomecircumstances.Plantpropagatorswanttomaintainjuvenilityin
ordertovegetativelypropagatecuttingswhileflowerandfruitgrowerswanttoreducethejuvenilephase.Earlierfloweringand
fruitingreducesproductioncostsandallowsforanearlierreturnoninvestments.

Environmentalfactorssuchasperiodsoflongorshortdaylight,varyingnutritionallevelsorsupplyingcarbondioxideenriched
atmospheremayincreasevegetativegrowthandifproperlycontrolledmayshortenthetimetomaturity.Theaffectthat
environmentalandhormonalfactorshaveonthelengthofjuvenilephasewilldependultimatelyongeneticcontrol(7).

Aplantisconsideredmaturewhenitbecomespotentiallycapableofreproducing.Althoughaplantmaybemature,floweringmay
notoccuruntilenvironmentalconditionsarefavorable(7).

ReproductiveGrowth

Thevegetativestageofgrowthendswhenthevegetativestemprimordiaistransformedintoflowerprimordia.Oncefloralinitiation
hasbeguntheprocessisirreversibleandwillcontinueevenifenvironmentalconditionswhichstimulatedinitiationhavechanged
(7).

Photoperiodismisthegrowthresponseofaplanttothelengthofthelightanddarkperiods.Shortdayplantsinitiatefloweringonly
whenthedaylengthislessthan12hours,andincludemanyspringandfallfloweringtemperateplants.Longdayplantsinitiate
floweringonlywhenthedaylengthisgreaterthan12hours,oraspecificcriticalperiod.Mostsummerfloweringplantsarelongday
plants.Dayneutralplantscaninitiatefloweringindependentofdaylength(7).

Thefloweringstimulusisformedinleavesandtransportedtotheapicalmeristeminresponsetothephotoperiod.Thetheorythat
afloweringhormoneflorigenisresponsibleforflowerinductionhasbeenpostulated,however,thissubstancehasnotyetbeen
isolated.Theleavesofsomeplantsinitiatefloweringinresponsetoonlyonecycleoftheproperdaylength.Mostplantsrequire
manycyclesofproperphotoperiodinordertoinitiatefloweringandmanysuchasthechrysanthemummayalsorequiretheproper
temperatureduringshortdaystoinitiateflowers(7).

Phytochromeisabluegreenpigmentfoundinallplants.ItisfoundintwoformsthePrformabsorbsredlight(660nm)andis
convertedintothePfrform.ThePfrformabsorbsfarredlight(730nm)andisconvertedbackintothePrform.Pfrisalsoslowly
convertedtoPrduringthedarkphase.ThenettransformationfromtheinactivePrformtotheactivePfrformduringthecourseof
achangingphotoperiodaffectsthefloweringmechanism(4).Interruptionofthedarkphase(nightbreak)byabriefperiodoflight
caninhibitfloweringofshortdayplantsandinitiatefloweringinlongdayplants.Redlight(incandescentlamps)arecommonly
usedtoeffectivelyproducenightbreak.Phytochromeisalsoresponsiblefortheinitiationandinhibitionofgerminationofsome
seeds,however,alightrequirementisnotnecessaryformostseeds(7).

Temperaturealsohasadirecteffectonflowering.Thetermvernalizationisusedtodenoteanycoldtemperaturetreatmenttoa
plantthatinducesflowering(4).Manybiennialsrequireaperiodoflowtemperaturetoinduceflowering.Aplantwhichhasbeen
givenacoldtreatmentcanbegraftedontoanonvernalizedplant,andbothwillflower(7).Thisimpliesthatasubstanceis
producedwhichpassesacrossthegraftuniontoinducefloweringinthenonvernalizedplant.Theflowerinitiatingsubstancehas
notyetbeenidentified,however,thecoldrequirementofsomeplantshasbeenreplacedbythemultipleapplicationofgibberellic
acid.

Althoughphotoperiodismandvernalizationareinterrelated,thestimuliproducedbyeacharenotidentical.Evenafteraplanthas
receivedthepropertemperatureforvernalization,floweringwillnotbeinitiateduntiltheplantisexposedtotheproperphotoperiod
(7).

Watermayalsoaffectflowerinitiation.Manyplantsshowmorefloweringinthespringwhentheprevioussummerandfallweredry
(7).

Fruitdevelopmentusuallyoccursconcomitantlywithflowering.Priortopollinationtheincreaseinfruitsizeisaresultofcell
division.Thestimuliandnutrientsforthisgrowtharesuppliedbytheplant.Pollinationisthetransferofpollenfromtheantherto
thestigmaandservestwofunctionsfirst,theinhibitionofflowerandfruitabscissionandsecond,toprovidethemalegametefor
fertilization.Thesetwofunctionsoccurseparatelyandeventhoughpollinationhasoccuredandfruitsetisobtained,fertilization
maynottakeplace.Thismaybeduetothefailureofthepollentogerminateorthepollentubetogrowfastenoughtoreachthe
ovarybeforeitisshed.Pollenrequiresthepresenceoforganicandinorganicsubstancesonthepistiltostimulateitsgermination.
Othersubstanceschemicallyattractthegrowthofthepollentubeandmaypreventfertilization.Fruitthatissetandgrowswithout

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fertilizationandthusdoesnotproduceviableseediscalledparthenocarpic.Seedlessfruitareoftenhorticulturallydesireable
howevervegetativepropagationmayberequiredtocontinuethecultivar(7).

Whenfertilizationoccurs,thedevelopingplantnolongerdependsontheparentplantforasourceofgrowthstimuli.Thestimuli
nowcomefromthedevelopingseed.Theeffectoftheseedonfruitdevelopmentischemicallymediated.Thegrowthregulators
auxin,gibberellinandcytokininplayaroleinfruitdevelopment.Theconcentrationofthesesubstancesvariesatdifferentstages
offruitdevelopmentandconsequentlyaffectfruitgrowthanddevelopment(7).

Foodmaterialsnecessaryforthedevelopingfruitaresuppliedbyvariousplantparts.Theavailabilityofnutrientsandmoisturewill
haveadirecteffectonfruitsize.Whenthenumberoffruitsetishigh,thesizeoftheindividualfruitswillbereduced.Removalof
someofthefruitatanearlystagewillallowtheremainingfruittoobtainmorenutrientsandwatertoproducelargerfruits.
Judiciousfruitthinningmayleadtolarger,betterqualityfruitandanincreasedprofitmargin.

Fruitripeningofmanybutnotallfruitscoincideswithaspecificphysiologicalprocess,theclimateric.Itischaracterizedbya
suddenriseintherespirationofthefruitresultinginaburstofcarbondioxideproduction.Pigmentchangesalsooccuratthistime
withpeakripenessoccurringatthepeakoftheclimatericorimmediatelythereafter(7).

Temperaturehasagreateffectontherateofmaturationandprogressoftheclimateric.Respirationratesincreasewithhigher
temperaturesincreasingcarbondioxideproductionanddecreasingfruitlife,particularlypostharveststoragelife(7).

Ethylene,agaseousplanthormone,isimportantinfruitripening.Ethyleneisproducedbythedevelopingfruit,byrottingtissue
andmaybeacontaminantofnaturalgas.Whenfruitisstoredunderconditionswhereethyleneisremovedfromtheenvironment,
ripeningcanbedelayed(7).

Senescencereferstotheprocessesinvolvedinthedeteriorationoftheplantoritsorganspriortodeath.Inannualsandbiennials,
senescenceoccursafterfloweringandfruiting.Inperrenialsitoccursasagradualdecreaseingrowthandviability.Senescence
canbepostponedinsomeplantshowever,deathisinevitable(7).

INTERNALFACTORSAFFECTINGPLANTGROWTH

GeneticThegeneticcomplimentofaplantisacquiredwhenthezygoteisformedfrommaleandfemalegametes(12).The
geneticinformationisduplicatedandpassedonwithsubsequentcelldivisions.Astheplantenlargestoitsmaturesizesome
genesareactivatedwhileothersareinactivated.Certaingenesdirectthesynthesisofenzymesthatcatalyzespecificbiochemical
reactionsrequiredforgrowthanddifferentiation.Thegenesinvolvedinproteinsynthesisarereferredtoasstructuralgenes.
Regulatoryandoperatorgenesregulatetheactivityofthestructuralgenes(4).Thesignalsthatstimulatetheregulatorygenesare
believedtobegrowthregulators,inorganicions,coenzymesandenvironmentalfactorssuchastemperatureandlight.

GrowthRegulators

Thetermhormonewasdevelopedbyanimalphysiologiststodenotenaturallyoccurringorganicsubstances,producedata
specificsite(usuallyagland),effectiveatlowconcentrations,whoseactionmaybeinvolvedatsitesfarremovedfromtheirorigin.
Thetermgrowthregulatorhasbeenusedtoincludeallnaturallyoccurringandsyntheticallyproducedsubstancesthataffectplant
growthanddevelopment(7).

Growthhormonesparticipateinbothgeneticandenvironmentalcontrolofgrowthanddifferentiation.Thepatternofdistributionof
growthhormonesintheplantiscontrolledbyinteractionsbetweentheenvironmentandgeneticfactorsintheplant(16).Theymay
beeithergrowthinhibitorsorpromotorsdependingonthesiteofactionandconcentrationofthesubstance.Thereare5major
typesofplanthormones:auxins,cytokinins,gibberellins,abscisicacidandethylene.

Auxinsaregrowthhormonesproducedinallhigherplants.Theyappeartobeformedinthemeristematictissuesofstemandroot
apices,youngdevelopingleaves,flowersandfruits(16).Thehighestrateofauxinbiosynthesisisintheshootapicalregion.Auxin
istransporteddownwardresultinginaconcentrationgradientinthevariousplantparts.Theresultantconcentrationofauxinhas
beencorrelatedtoinhibitionandstimulationofgrowthaswellasdifferentiationoforgansandtissues(7).

Auxinsinfluenceplantgrowthinmanywaysincludingcellenlargementandelongation,phototropism,geotropism,apical
dominance,abscissionofplantparts,flowerinitiationanddevelopment,rootinitiation,fruitsetandgrowth,tuberandbulb
formation,andseedgermination.Commerciallysyntheticauxinsareusedtoinitiateadventitousrootsfromcuttings.Indolebutyric
acid,indoleproprionicacidandnaphthaleneaceticacidaresynthecticauxinsappliedtothebasesofstemcuttingstostimulatethe
initiationofadventitousroots(4).

Weedcontrolbyanothersyntheticauxin,2,4dichlorophenoxyaceticacid(2,4D),iswidespreadasaselectiveherbicideagainst
broadleefweeds.Auxinsarealsousedtoincreasefruitset.Useof4chlorophenoxyaceticacidtoincreaseblossomandfruitset
intomatoesisalsosuccessful.Auxinsarealsocommonlyusedintissuecultureprocedurestoinitiaterootinginexplantsorcallus
(4).

Gibberellinsareagroupofnaturallyoccurringplanthormonesthataffectcellenlargementanddivisionwhichleadstointernode
elongationinstems.Theyhaveadwarfreversingresponseallowingcertaindwarfcultivarstogrowtonormalheightwhentreated
withgibberellin.Theyaffectmanydevelopmentalprocesses,particularlythosecontrolledbytemperatureandlightsuchasseed
andplantdormancy,germination,seedstalkandfruitdevelopment(7).

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Gibberellinsareusedcommerciallytoincreasefruitsizeof"ThompsonSeedless"grapes.Theyareappliedatfruitsetorshortly
thereafter.Theyalsopromotemaleflowerinitiationincucumberswhenpolleniswantedforhybridseedproductionandmay
overcomethecoldrequirementforfloweringofsomeperennialplants(4).

CvtokininsDrimarilvDromotecelldivisionbuttheyalsoinfluencecellenlargement,tissuedifferentiation,dormancy,phasesof
floweringandfruitingandretardationofleafsenescence(4).

Cytokininsandauxinsinteracttoaffectdifferentiation.Ahighauxintolowcytokininratiostimulatesrootdevelopment,whereasa
lowauxinandhighcytokininratiostimulatesbuddevelopment.Equalconcentrationsofauxinandcytokininresultsin
undifferentiatedtissueorcallus(7).

Cytokininsarenotcommonlyusedinagriculture,however,cytokininmaybeusedintissueculturetoinduceshootdevelopment
(4).

Ethyleneisagasthatdiffusesreadilythroughouttheplant.Itisproducedinmeristematictissues,ripeningfruits,senescing
flowersandfruitsandgerminatingseeds.Thecuticularcoatingoftheplanttendstopreventlossesfromtheplant(4).

Syntheticethylenereleasingcompoundssuchasethephonhaveseveralvaluablecommercialapplications.Ethephonisusedto
ripenbananas,pineapples,melonsandtomatoes,andwhenappliedasapreharvestsprayitpromotesuniformripeningofapples,
cherriesandpineapple.Itisusedtoincreasetheproductionoffemaleflowersoncocumberswhichdevelopfruitsandincrease
yields.Highconcentrationsofethylenemaybeharmfultoplants,inducingleafabscissionandhasteningsenescenceofflowers
andfruits(4).

Abscisicacidinteractswithotherhormonesintheplant,counteractingtheirgrowthpromotingeffects.Itinhibitsratherthan
stimulatesplantgrowth.Abscisicacidpromotesdormancyinseedsandisinvolvedinleafandfruitabscision.Theabscisicacid
contentofleavesincreasesfollowingwaterstress,whereitinducesclosureofthestomata(4).Abscisicacidisexpensiveto
synthesizeandnocommercialapplicationsareasyetinuse.

Greenhousegrowersandnurserymencommonlyusegrowthretardantsinmanagingplantgrowth.Manysyntheticcompoundsare
availabletodwarfplants,increasebranchingandmanagefloweringtoproducecompactfloweringplantsinatimelymanner.Use
ofgrowthretardantsisspecificbyspeciesanddesiredresult.

EXTERNALFACTORSAFFECTINGPLANTGROWTH

Plantgrowthanddevelopmentareinfluencedbyphysical,chemicalandbiologicalcomponentsintheplantsenvironment.Any
factorintheplants'environmentthatislessthanoptimum,whetheritisdeficientorinexcess,willlimitplantgrowth(17).

Light

Plantsrespondtolightofthewavelengthsfrom300800nm.Plantsgrownintheabsenceoflightaresaidtobeetiolated.Etiolated
plantslackchlorophyll,aretallandspindlywithlonginternodesandhavesmallleavesthathavefailedtoexpand(12).Their
morphologicalexpressionofetiolationisrelatedtotheeffectoflightonauxindistributionandsynthesis(4).Thereareno
anatomicaldifferencesinthetissuesformedinthelightordark,however,lightacceleratesmanyphasesofgrowthwhileinhibiting
certainaspectsofinternodeelongation(12).

Lightcanhaveaneffectonthemorphologyoftheplant.Leavesonthesameplantmaydifferdependingonwhethertheyaresun
leavesorshadeleaves.Sunleavesareoftenthickerwithextralayersofpallisadeparenchyma,andshorterpetioles.Theyare
alsosmallerinarea(12).

Aplant'sresponsetolightwillvarydependingontheintensity,durationandwavelengthofthelightitreceives.

Lightintensityreferstotheconcentrationoflightwavesstrikingtheleafsurface(7).Lightintensityhasbeenexpressedin
footcandlesbyscientistsandgrowersuntilrecently.Wattspersquaremeterormicroeinstein'spersquarecentimeteraremore
usefulanddescribeenergyperunitareawhichcanberelateddirectlytopowerconsumptionforcostanalysis.

Lightintensityishighwheretherearenocloudsandlittlemoistureintheair.Watervaporintheatmosphereabsorbsradiationso
lightintensityislowerincloudyorhumidareas.Lightintensitywillvarywiththeelevation,latitude,seasonandtheweather
conditionsaffectingtheamountofwatervaporintheair(4).

Photoprocessesintheplantvaryintheintensityofthelightrequiredtoinitiatethereactionsandtheeffectoftheintensityonthe
rateofthereaction(7).Therateofphotosynthesisdropsoncloudydays.However,notallplantsrequirehighlightintensities.
Shadeplantsmayrequireaslittleas1/10fullsunlightforoptimumgrowthandhigherlevelsmaycausesunburning,scaldandin
severecasesdeathifsufficientsoilmoistureisnotavailable.

Photoperiodismreferstothephysiologicalresponsesofplantstovariationsinthedurationofdaylight(4).Theshiftfrom
vegetativegrowthtoreproductivegrowthisaresponsetothephotoperiod.Thelengthofthevegetativegrowthperiodcanbe
extendedbygrowingplantsinphotoperiodsthatdonotinduceflowering.Daylengthmayalsoaffectthetimetofirstflower,the
numberofflowersproducedandthenumberoffruitset(12).Likewiseshortdaysandcoolertemperaturesinitiatedormancyin
manytemperatezoneperennialplants.

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Thelightreactionsoftheplantarecarriedonbydifferentpigmentsystemsthatabsorbspecificwavelengthsoflight,i.e.,blue,
green,yelloworredlight(12).Chlorophyllabsorbsthatradiantenergynecessaryforthephotoprocessesofphotosynthesis(7).
Chlorophyllabsorbslightintheredandblueportionsofthespectrum(7).Phototropism,themovementorbendingofstems,
leavesandflowerstowardlight,istriggeredbybluelight(4).Thisprocessisbelievedtooccurduetotheaccumulationofauxinin
theshadedsidepromotingcellgrowth.Thusthebendingmovementtowardthelightsourceisaresultofincreasedcellgrowthon
theshadedside(4).Whenleavesaresubjectedtohighlevelsofradiation,theymayorientthemselvesparalleltotheenergy
sourceinordertominimizetheharmfuleffectsoftheintenseradiation(12).

Althoughincominglightinthetypicalgreenhouseinmidwinteroftendoesnotexceed10001500footcandlesinmanylocations,
goodgrowthoflettucemaybeobtainedatintensitiesaslowas500footcandles.Boltingresultsfromlongdaysandhightempera
turessomostvarietiesofgreenhouselettucearenotgrowninlatespringandearlysummer(18).

Earlyspringcucumbers,attheseedlingstage,respondtosupplementallight.Daylengthofabout1214hourswith18002000foot
candlesattheplantlevelshouldbeprovided.Crowdingshouldbeavoidedtopreventplantsfrombecomingspindly(18).

Tomatoesgrowninthelatefallorearlywintershouldbeexposedtoasmuchlightaspossibleduringnormaldaylighthours.
However,artificiallightsshouldnotbeusedtoextendthedaylengthastomatoesareplantswhichflowerandfruitbetterifdayl
engthistwelvehoursorless.Ifartificiallightsareused,atleast500footcandlesattheleafsurfaceshouldbeprovided.
Supplementaryartificiallightmayonlybeeconomicallyfeasiblefortomatoesattheseedlingstagewhereagreaternumberofp
lantscanbeilluminatedpersquarefoot(18).

Temperature

Thetemperaturerangethatsupportsplantgrowthisgenerallyfrom4097degreesF(4.536degreesC)(7).Optimumtemperatures
forgrowthvarywiththespeciesandthestageofdevelopmentandusuallyfluctuatesnighttoday.

Severalgrowthprocessesaresensitivetotemperature.Amongthesearerespiration,partofthephotosyntheticprocess,
maturation,flowering,fruitripeninganddormancy(7).

Photosyntheticratesaredeterminedmainlybylightintensity,CO2levelsandtemperature(11).Temperaturehaslittleeffecton
photosyntheticratefrom5086degreesF(1530degreesC)untillightandCO2becomesaturatedforthephotosyntheticprocess.
Atthispoint,anincreaseintemperaturefrom6886degreesF(2030degreesC)resultsinamarkedincreaseinthe
photosyntheticrate(11).Onwarmdays,middayleaftemperaturesmaybehighandinhibitphotosytheticactivity(12).Notonlyare
metabolicprocessesreducedathighleaftemperatures,butmoisturestress,fromincreasedtranspirationallosses,resultsin
stomatalclosurewhichdecreasesthesupplyofCO2tothechloroplastsslowingphotosynthesis.

Respirationratesincreaserapidlyasthetemperatureincreases.Temperatureisacontrollingfactorinestablishingthe
compensationpointofgreenhousecrops,thepointatwhichtherateofCO2consumedinphotosynthesisequalstherateofCO2
givenoffinrespiration,becauseofitsaffectonrespirationrate(11).Astemperaturesrisethelevelatwhichthecompensation
pointoccursforaparticularlightlevelorCO2concentrationwilldecrease.Acessationofgrowthoccurswhentherateof
respirationincreasesmorerapidlythantherateofphotosynthesis,resultinginadepletionoffoodreserves(7).

Maintainingday/nighttemperaturesatspecificlevelscanincreaseyieldandqualityofcrops.Optimumgrowthofmanycrops
occurswhengreenhousetemperaturesarecooleratnightthanduringtheday.Theresponseofplantstodiurnaltemperature
fluctuationsisreferredtoasthermoperiodicity(11).

Temperatureeffectsonfloweringmaybedirectorinductive(11).Theeffectoftemperatureisdirectwhenflowerinitiationoccurs
duringtheperiodoftemperaturetreatment.Ifaspecifictemperatureinducesachangewithintheplantwhichpermitsfloweringat
anothertime,theeffectisconsideredtobeinductive.Vernalizationistheinductiveeffectofcoldtemperaturesonflowerinitiation.
Manybiennialsandperennialsrequirecoldtreatmentstoinduceflowering.

Roottemperaturesalsoaffecttherateofplantgrowth.Increasingroottemperaturesuptoabout26degreesC(76degreesF)may
increasetopgrowthandtheuptakeofinorganicions.Thisistrueofmanyhydroponicallygrowncrops,cucumbersinparticular(2).

Gases

Greenplantsrequireoxygenfornormalgrowthanddevelopment.Theenergyreleasedincellularrespiration,fromthebreakdown
ofcarbohydratesandcomplexorganicmolecules,consumesoxygenandreleasesCO2.Mostplantsrespirecontinuously,day
andnight,requiringacontinuoussupplyofoxygen.Anaerobicrespirationorfermentationoccursintheabsenceofoxygen.The
productsofthisformofrespirationareoftendeleterioustotheplantandtheenergyreleasedisrelativelysmallcomparedto
aerobicrespiration.Rootsalsorequireoxygenforaerobicrespirationwhichtheyobtaindirectlyfromthegrowingmedia.The
absorptionofsaltsandrootextensionaredependentupontheenergysuppliedfromrespiration.Poorlyaeratedgrowingmedias
resultinadecreaseinwaterabsorptionduetoareductioninthepermeabilityoftherootcells.Afterextendedperiodsofpoorroot
aerationtherootsstopgrowingandaremoresusceptibletodisease(11).

Seedsrequireoxygentogerminate.Seedgerminationisinhibitedbyalackofoxygenforprolongedperiods.Oftenthickoroily
seedcoatsmustberemovedfromtheseedsooxygenwillbeavailabletotheembryo.Compactedorwaterloggedsoilsor
growingmediacanalsocreateanoxygenlessenvironmentandseedswillnotgerminate.

CarbonDioxide

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Carbondioxide(CO2)isarawmaterialrequiredforphotosynthesis.TheatmosphericCO2concentrationattheplantlevelisthe
mostimportantratedeterminingfactorforfurtherincreasesinphotosynthesisandyield(18).CO2concentrationsmayfallbelow
theambientairconcentration.03%(300ppm)inthegreenhousewhenweatherconditionsrestrictventilationorinfiltration.Acrop
inatightlyclosedgreenhousewillsoondepletetheCO2concentrationwhichreducesgrowthandproductionbyslowingor
stoppingphotosynthesis.Unlessreplaced,theCO2concentrationwillremainattheplantscompensationpoint,thelevelatwhich
theCO2producedfromrespirationequalstheamountusedforphotosynthesis.Nogrowthoccursatthispoint.

Whenweatherconditionspermit,ventilationisaneffectivemethodofmaintainingCO2concentrationsatthenormalairlevels.
However,plantsrespondfavorablytohigherCO2concentrations,makinggreenhousesupplementationofCO2aneffective
methodofincreasingplantgrowth(11).AlthoughtheCO2responseisdependentuponlightintensity,beneficialeffectsare
obtainedoverawiderangeoflightintensities,eithernaturalorartificial.CO2enrichmentisofspecialsignificanceinhydroponic
cultureasdecayingorganicmatterinthesoil,asourceofCO2,isnotpresent(18).

CO2iscommonlysuppliedat8001600ppmviagasCO2generatorsorlargetanksofliquidCO2dependinguponthecost
comparisonbetweenthetwoandtheavailabilityofthebottledcarbondioxide.

Airpollutants

Airpollutionisanimportantproblemforproducersofgreenhousecrops.Thesourcesofairpollutionareincreasingasnew
industriesandhighwaysarebuilt.Thisisaparticularproblemforhorticulturaloperationsnearurbanandindustrialareas.Among
thephytotoxicpollutantsareozone,peroxyacelnitrates,oxidesofsulfur,hydrocarbons,fluorides,carbonmonoxide,herbicides,
fumigants,mercuryvapors(donotusemercurythermometersingreenhouses),andphytotoxicgasesproducedfromincomplete
combustionofCO2generators(7).Itmaybenecessaryforgreenhouseownerstomovetoareaswherephytotoxicgasesarenot
present,ortogrowspeciesthatarelesssensitivetothesesubstances(11).

Oftenleavesandflowersarefirsttoshowsignsofairpollution.Unusualdiscolorations,spotting,twistingorturningofleavesand
abortionofflowersfollowedbypoorgrowtharesymptomsofairpollution.

Water

Mostgrowingplantscontainabout90%water(4).Wateristhemediumfortransferwithintheplantandisthesolventsystemof
thecell.Waterisoneoftherawmaterialsforphotosynthesisrequiredfortheproductionofnewcompounds.Insofttissueswater
pressureprovidessupportandasplantslosewaterfromtheirleavestheyarecooled(7).Anetlossofwaterwillcausegrowthto
stopandcontinueddeficiencyresultsindeath.

Agrowingplantabsorbswaterfromthesoilandgivesitoffintranspiration.CO2enterstheplantthroughafilmofwaterthat
surroundstheleafandasthefilmevaporatesitisreplenishedbytheplant.ThetranspirationallossofwaterinexchangeforCO2
isnecessaryforplantgrowth.Rapidlygrowingplantsrequirelargequantitiesofwater,farinexcessofthatfoundintheplantfor
synthesisofnewmaterials(7).

Moisturestressisgenerallydetrimentaltoplantgrowthreducingbothyieldandqualityofthecrop.Thedegreeanddurationofthe
stresswilldeterminehowseverelygrowthisreduced,however,growthratemayneverreturntothelevelitwasbeforethestress
(11).

Thestageofgrowthwhenmoisturestressoccursisalsoimportant.Moisturestressatthetimeofflowerinitiationmay
significantlyreduceyield.Severestressleadstoprematureflower,leafandfruitdrop(11).

Transpirationleadstomoisturestressifmoistureisnotreadilyavailabletotheroots.Asmoisturestressincreases,stomates
closeandphotosynthesisisreduced.Warmdryairhasahighevaporativecapacity,increasingtherateoftranspiration.Aswell,
theincreaseinleaftemperatureresultingfromhighlightintensityraisestherateoftranspirationalloss(11).

Poorwaterqualitycanbeamajorproblemforgrowers,particularythosewithhydroponicsystems,duetocontaminationfrom
organicandinorganicsubstances.Eventhebestdomesticwatersuppliesmaycontainsubstancesthataffectplantgrowth.
Therefore,acompletewateranalysisisrecommendedforgreenhousegrowers.Hydroponicsystemsrequiredetailedelemental
analysisofirrigationwaters.Inordertodevelopanappropriaterecommendationfornutrientlevelsinsolutiontheconcentrationof
existingelementsinthewatermustbeknown.Adjustmentscanthenbemadeinthesolutionforthecroptobegrown.Depending
ontheresultofthewateranalysis,someformofwatertreatmentmaybenecessary.Watertreatmentmaysimplyinvolvetheuse
ofafilteringsystemforparticulatedebris,ormayrequiremoresophisticatedmethodsofionexchangeorreverseosmosisin
additiontofiltration.Insomecasesallthatmaybenecessaryistheadjustmentofnutrientsolution,asinhardwaterareaswhere
themajorityofcalciumandmagnesiumisalreadyprovidedbythewatersource(9).

Nutrition

Sixteenelementsareconsideredtobeessentialforgrowthanddevelopmentinhigherplants.Arnon&Stout(1)consideredan
elementessentialwhenit1)isrequiredbyaplanttocompleteitslifecycle,2)theactionoftheelementisspecificandnoother
elementmaybesubstitutedforitand3)theelementmustexertitseffectdirectlyongrowthormetabolismandnotsimplycause
anotherelementtobemorereadilyavailableorantagonizeatoxiceffectofanotherelement.

Theessentialelementsaredividedintotwogroups:themacronutrients,thoserequiredinrelativelylargequantitiesincluding
carbon,hydrogen,oxygen,nitrogen,phosphorus,potassium,calcium,magnesiumandsulfurandthemicronutrients,those

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requiredinsmallquantities,includingiron,chlorine,manganese,boron,zinc,copperandmolybdenum(SeeTable1forInternal
ConcentrationsofEssentialsElementsinPlants).

Carbon,oxygenandhydrogenareobtainedfromtheenvironment,specificallycarbondioxideorwater.Alongwithchlorine,which
isfoundinmostwatersources,theseelementsaregenerallynotconsideredintheformulationofnutrientsolutions.

Table1.InternalConcentrationsofEssentialElementsinHigherPlants
ConcentrationinDryTissue

%
Element ppm

Hydrogen 60,000 6

Carbon 450,000 45

Oxygen 450,000 45

Nitrogen 15,000 1.5

Potassium 10,000 1.0

Calcium 5,000 0.5

Magnesium 2,000 0.2

Phosphorus 2,000 0.2

Sulfur 1,000 0.1

Chlorine 100 0.01

Boron 20 0.002

Iron 100 0.01

Manganese 50 0.005

Zinc 20 0.002

Copper 6 0.0006

Molybdenum 0.1 0.00001

AdaptedfromSalisburyF.B.andRossC.1969.PlantPhysiology.Belmont,Calif.Wadsworth,p.82.

FunctionoftheEssentialElementsinPlants

Nitrogen

Nitrogenisaconstituentofaminoacids,proteins,coenzymes,nucleicacidsandchlorophyll.Nitrogenhasagreataffectonplant
growthandadeficiencyorexcessmarkedlyaffectsplantgrowthandfruityield(3,8,9,13).

Nitrogenisamobileelementintheplantanddeficiencysymptomswilldevelopfirstonlowerleavesasthenitrogenisremovedfor
useinnewdevelopingleaves.Theolderleavesbecomechlorotic(turnyellow)andeventuallydie.Nitrogendeficiencycanhavea
considerableeffectonfinalyieldifitoccursforprolongedperiodsduringcriticalstagesofgrowth(3,8,9,13).

Toomuchnitrogenproduceslushplantswithdarkgreenthinfoliagewithfewblossomsandfruitsetisadverselyaffected(3,9).

Inhydroponicsystemspropercontrolofnitrogenconcentrationaswellastheformoftheelementisimportant.Plantstakeupthe
nitrate(NO31)andammonical(NH4+1)formsofnitrogen.Aproperbalancebetweenthenitrateformandammoniumformis
requiredforplantgrowthandalsoprovidessomedegreeofpHcontrol.Aratioof75%nitrateto25%ammoniumissatisfactoryfor
nutrientsolutionsandshouldnotexceedaratioof50/50orammoniumtoxicitymayresult(3,9).

Mostnutrientsolutionformulascallfor100200ppmnitrogenwithadesiredratioofnitratetoammoniumionsat3or4to1.Some
solutionsmaystartthenitrogenatalowerleveltominimizevegetativegrowthandpromotefruitinitiationanddevelopment(9).

Nitrogenisakeyessentialelementaffectingplantgrowthandcropyields.Successinhydroponicgrowingsystemsmaydepend
toalargeextentonthemanagementofthiselement(9).

Phosphorus

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PhosphorusisaconstituentofATP,nucleicacids,phospholipidsandcertaincoenzymes.Itisveryimportantintheplantsenergy
transfersystemandadeficiencycanslowgrowthconsiderably(3,8,9,13).

Phosphorusoverfertilizationmaybeaproblemwithsoillessculture.Phosphorustoxicitymayoccur,interferingwiththenormal
functionofotherelementssuchasiron,manganeseandzinc(3,8,9).

Phosphorusdeficiencyreducesgrowthandolderleavesdevelopapurplishcolorasanthoyaninpigmentsaccumulate.Phosphorus
uptakeisinfluencedbytemperatureandadeficiencymaybeinducedbycoolnutrientsolutiontemperatures(3,8,9,13).

Mostformulascallfor3050ppmofphosphorusintheformofmonoorallhydrogenphosphateanions(HPO4orH2PO41)oras
phosphoricacid(H3PO4)

Potassium

Potassiumactsasacoenzymeoractivatorofmanyenzymesystems.Highpotassiumlevelsarerequiredforproteinsynthesis
andfruitproductioninmostcrops(i.e.,tomato)asthedemandforpotassiumbythedevelopingfruitishigh.Adeficiencyduring
fruitingproducesfruitofsignificantlylowerqualityandsize(3,8,9,13).

Potassiumdeficiencysymptomsbeginasslowgrowth.Ifthedeficiencybecomesseverelowerleavesdevelopamarginal
chlorosisgivingtheappearanceofburnededges(3,8,9,13).

Acriticalbalanceisrequiredbetweenpotassium,calciumandmagnesiumorplantstressoccurswhenthelevelofpotassiumis
highincomparisontocalciumormagnesium.Highlevelsofpotassiuminsolutionmayinduceacalciumormagnesium
deficiency.Caremustbetakentomaintaintheproperbalancebetweenthesethreeelements.Mostnutrientsolutionscallfor200
ppmpotassiuminsolution(9).

Calcium

Calciumisrequiredtomaintainmembraneintegrityandisfoundincellwallsascalciumpectatewhichcementstogetheradjacent
cellwalls(13).

Calciumdeficiencyisgenerallyaresultofanimbalancewithpotassiumandmagnesium.Itprimarilyaffectsleafsizeandshape
andisthecauseofblossomendrotindevelopingfruit(3,8,9,13).

Aconcentrationof200ppmofcalciumiscommonformostnutrientsolutionformulas.Sincecalciumiscommoninmanynatural
watersources,awateranalysisisnecessarysoadjustmentscanbemadeinordertoavoidoverfertilizationwhichwouldleadto
animbalancewithpotassiumandmagnesium(3,8,9,13).

Magnesium

Magnesiumisanessentialconstituentofchlorophyllandisrequiredforactivationofmanyenzymesinvolvedintheenergy
transferprocesses.Adeficiencyofmagnesiumwillseriouslyaffectplantgrowthanddevelopmentasphotosynthesisisdirectly
affected(3,8,9,13).

Deficiencyofmagnesiumfrequentlyoccursduetoanimbalancewithpotassiumorammoniumions,appearingasinterveinal
chlorosisdevelopingfirstonolderleaves.Magnesiumexcessesrarelyoccurhowever,magnesiumconcentrationsshouldnot
exceedthatofcalciumtomaintainapropercationbalance(3,9).

Theconcentrationofmagnesiumcalledforinmostnutrientsolutionformulasis50ppm.Irrigationwatersmaycontainhighlevels
ofmagnesiumawateranalysisisnecessarytomanagethelevelproperly(9).

Sulfur

Sulfurisaconstituentofsomeaminoacidsandproteins,coenzymeA,thiamineandbiotin(3,9,13).

Theratioofsulfurtonitrogenmaybeabettermeasureofthesufficiencyofsulfurintheplantratherthantotalsulfur
concentration.Deficiencysymptomsappearsimilartonitrogendeficiencysymptomsasanoverallplantyellowingorchlorosis.
However,sulfurdeficiencysymptomsstartinnewleaves(becauseitisnottranslocatedoutofoldleaves)wherenitrogen
deficiencysymptoms,appearfirstinolderleavesasnitrogeniseasilytranslocatedtonewleaves.Aplantanalysismaybe
necessarytodeterminewhichelementisdeficientiftheoverallplantischlorotic(3,9,13).

Mostnutrientsolutionformulascallforapproximately50ppmsulfurintheformofthesulfateanion(SO42).Highconcentrationof
SO42ionsgenerallydonotcauseanyharm(9).

Boron

Theroleofboroninplantsisnotwellunderstoodalthoughthereisevidencethatitisimportantincarbohydratesynthesisand
transport.Minutequantities(<.5ppm)areusuallyrequiredbyplants,andmanyaresensitivetohigherlevelsofthiselement(3,8,
13).

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Borondeficiencywillslowgrowthoftenstuntingthewholeplant.Fruitdevelopmentwillbeslowandfruitqualitypoor(3,9).

Borontoxicityfromexcessboroninthenutrientsolutionorboroninthewatersupplyresultsindiscolorationandeventualdeathof
theleafmargins(3,8,9,13).

Nutrientsolutionformulasusuallycallforabout.3ppmboroncommonlyintheformofborateanion(BO33)orboricacid(H3BO3).
WellwaterinthewesternU.S.maycontaintoxicorneartoxiclevelsofboron.Testirrigationwaterstoknow(3,9,13).

Chlorine

Chlorineisrequiredinphotosynthesisasanenzymeactivatorduringtheproductionofoxygenfromwater(13).

Chlorineisrarelyeverdeficientasitisacommonconstituentinwaterandchemicalsusedtopreparenutrientsolutions.Itdoes
notnormallyhavetobeaddedtothenutritionprogram.Anexcessofchlorinewillresultinburningofleafmarginsandpremature
lossofleaves(9).

Copper

Copperactsasanelectroncarrierandasaconstituentofcertainproteinsandenzymes.Acopperdeficiencyresultsinplantsthat
arestuntedwithchloroticolderleaves,whiledevelopingfruitaresmallandpoorlyformed(3,8,9,13).

Hydroponicnutrientsolutionsrequirefrom0.01to.1ppmcopper,foundinthenutrientsolutionasthecupriccation(Cu+2).In
hydroponicgrowingsystems,ifthecopperconcentrationgetstoohighrootdamagemayresult(3,9).

Iron

Ironisrequiredforthesynthesisofchlorophyllandisanessentialpartofthecytochromeswhichserveaselectroncarriersin
photosynthesisandrespiration(3,9,13).

Irondeficiencyappearsasanintervernalchlorosis,oftheyoungerplanttissue,whichmaybeconfusedwithotherelemental
deficiencies.Aplantanalysismaybenecessarytodeterminetheactualcause(9).

Ironeasilycomplexeswithmanysubstancessotheuseofachelatedformofironisgenerallycalledforwhenmakingnutrient
solutions,otherwiseitwillcombinewithotherelementsandbecomeaninsolubleprecipitateinthebottomofthesolutiontank.A
concentrationof23ppmofironineithertheferric(Fe+3)ortheferrous(Fe+2)formmustbemaintainedtopreventirondeficiency
(3,9,13).

Manganese

Manganeseactivatessomeoftheenzymesinvolvedinfattyacidsynthesis,DNAandRNAformationandtheenzymeisocitrate
dehydrogenaseintheKrebscycle.Itisinvolvedinproductionofoxygenfromwaterinphotosynthesisandmaybeinvolvedin
chlorophyllsynthesis(3,8,9,13).

Manganesedeficiencyappearsasinterveinalchlorosisontheyoungeplanttissueandmayresultinsignificantreductioningrowth
ifsevereManganesetoxicityappearssimilartodeficiencysymptomsatfirstthenbrowspotsonolderplanttissueandblack
specksonstemsandfruitdevelo(3,8,9,13).

Nutrientsolutionscallfor0.5ppmmanganeseintheformofmanganessulfate(3,9,13).

Molybdenum

Molybdenumisrequiredfornitrogenfixationbysymbioticnitrogenfixinbacteriaandforthereductionofnitratebytheenzyme
nitratereductas(3,9).

Molybdenumdeficiencymayappearasnitrogendeficiencyandresultsirestrictedplantgrowthandflowerdevelopment.Flower
abortioniscommowithmolybdenumdeficiency.Aplantanalysismaybenecessarytodetermincauseofthedeficiency(3,8,9,
13).

Nutrientsolutionscallfor0.05ppmmolybdenumwithammoniummolybdatecommonsource(9).

Zinc

Zincisrequiredfortheformationofthehormoneindoleaceticacidanisanenzymeactivator(3,8,9,13).

Zincdeficiencyresultsinstuntingofplantandleafgrowthandwhensevereleavesdieandfalloff.Deficiencysymptomsmaybe
confusedwiththdeficienciesofmagnesium,ironandmanganeserequiringaplantanalysistodeterminewhichelementis
deficient(3,8,9,13).

Nutrientsolutionformulascallfor0.05ppmofzincwhichiscommonlyappliedintheformofzincsulfate(9).

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FACTORSAFFECTINGNUTRIENTUPTAKE

Nutrientsaregenerallyabsorbedagainstconcentrationgradientconsequentlyrespiratoryenergyisrequiredfornutrientuptake(9).
Inorderforrespirationtocontinueintheroots,oxygenmustbeavailableintherootzone.Rootswhichbecometotallysubmerged
orwaterloggedforlongperiodwillsufferfromalackofoxygen.Thisleadstoslowgrowth,senescencearabscissionofleavesand
adventitiousrootingofstems(6).

Thesolubilityofoxygeninwaterislowanddecreasesfurtherasthsolutiontemperatureincreases.Increasesinroottemperature
(uptoabout30degreesC)increasesrespirationrateoftheroots,furtherincreasingoxygendemand,requiringaconstant
replenishmentofoxygentothenutrientsolution(9).

Rootzonewarmingformosthydroponicsystemsiseasilymanipulatedtosuitthecrop.Warmerroottemperaturesincreasesthe
rateofgrowth(2)andtheabsorptionandutilizationofnutrients(8).Temperaturealsoinfluencesthegrowthanddevelopmentof
rootsaffectingtheamountofrootsurfaceavailablefornutrientandwateruptake(9).

ThepHofthenutrientsolutionaffectstheavailabilityofelements.TheidealnutrientsolutionpHisbetween6.0and6.5.ApH
below5.0orabove7.0mayadverselyaffectplantgrowthbyalteringselectednutrientavailability.Themicronutrientsarep
articularlyaffectedwithexcessiveuptakeatlowpHsandremovalfromthesolutionthroughprecipitationathigherpHs(9).

Salinewater(watercontainingsodiumchloride)canbeusedinhydroponicsystemsifmoderatelysalttolerantcropssuchas
carnations,tomatoes,cucumbersandlettucearegrown.Salinewatershaveanhighosmoticpressurewhichreducesthewater
uptakebytherootsresultingininhibitionofplantgrowth.Ayieldreductionof1025%canbeexpected,dependingonthespecies,
varietyandsalinityofthewater(13).

Salinewateralsoreducestheavailabilityofcertainmicronutrients,especiallyiron,requiringadditionalirontobeadded(13).

Thetotalsaltconcentration,measuredbyelectricalconductivity(EC)isusedtomonitorthestatusofthenutrientsolution.Most
plantsgrowwellinanECofbetween1.8and3.5mmhos.TheECfallsastheplantsabsorbnutrientsfromthesolution.However,
theECdoesnotmeasurewhichnutrientsarebeingdepleted,andwithtimeselectedelementsmayaccumulateastheyarenot
removedquicklybytheplants.AdeficiencyoftheseusedinlargequantitiesmaybecreatedbutnotdeterminedbytheEC
readingwhichismaskedbyaccumulatedions.Toxiclevelsofcertainelementsmaydeveloprequiringthesystemtobeflushed
andanewsolutionmadeup(5).

Nutrientsolutionsarebeingdevelopedtoovercometheproblemsencounteredbytheadditionoflowdemandnutrientionsinthe
formulationofnutrientsolutions.Magnesiumandpotassiumsulfatescreateabuildupoflowdemandnutrientionsinrelatively
largequantities.Useofpotassiumandmagnesiumnitrateandtheadditionofmicronutrientsinchelatedformsreducethisbuild
up.Aswell,chelatedmicronutrientsareavailabletoplantsoverawiderrangeofsolutionpH.Studiesperformedonlettuce
demonstratedthisproblem.TheuseofanECmetertomonitornutrientsolutionswasnotsatisfactory.AlthoughtheECremained
inrecommendedranges,potassiumandnitratelevelswereverylowwithintwoweeks,resultingindeficiencysymptomsinthe
lettuceplants.Solutionsweredesignedsothationswereaddedtothenutrientsolutioninratiossimilartotherateofutilizationby
theplant,maintainingastablecompositionforlongerperiodsoftime.Growthwasvastlyimproved(15).

Moreworkisnecessarytodevelopnutrientsolutionsadaptedforgrowthofcucumbers,andtomatoesandselectedgreenhouse
crops.Careshouldbetakentomonitortheindividualelementcontentofnutrientsolutionstopreventthebuildupoflowuse
elementsandtheimbalanceddepletionofhighlyselectednutrients.Nutrientcompositionshouldmatchtheneedsofthecropas
growthproceedsattherateimposedbythemostlimitingfactorwhetheritbedeficientorinexcess(17).

Table2.NutrientConcentrationsinLeavesonaDryWeightBasis

Nutrient Tomatoes Cucumbers Lettuce

NormalRange NormalRange NormalRange

Nitrogen(%)

TotalN 3.05.0 2.55.0 2.15.6

Nitrate 1.21.5 0.81.8 2.59.3

Phosphorus(%) 0.40.8 0.51.0 0.50.9

Potassium(%) 4.08.0 3.06.0 4.010.0

Calcium(%) 1.54.0 2.08.0 0.92.0

Magnesium(%) 0.41.0 0.40.8 0.40.8

Sulfur(%)

TotalS 1.03.0 0.40.8 0.20.5

Boron(ppm) 2060 4060 2565

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Iron(ppm) 50150 90150 50500

Manganese(ppm) 2550 50150 25200

Copper(ppm) 46 410 518

Zinc(ppm) 1525 4050 30200

Molybdenum(ppm) 15 13 0.53

AdaptedfromHydroponicsWorld:StateoftheArtinSoillessCropProduction,AdamJ.SavagePh.D.,Editor,andKnotts'
HandbookForVegetableGrowers

Table3.NutrientConcentrationsforTomatoesinNFT

NO3 150200

NH4 0200

K 300500

P 50

Ca 150300

Mg 50

Fe 3

Mn 1

Cu 0.1

Zn 0.1

B 0.30.5

Mo 0.5

AsadaptedfromTheNutrientFilmTechniqueHorticulturalReview,ChrisJ.Graves.

Table4.NutrientConcentrationforLettuceinGrodanRockwool

Nutrient Concentration(ppm)

Nitrate 200

Phosphorus 60

Potassium 300

Calcium 170

Magnesium 5.0

Iron 3.0

Copper 0.1

Boron 0.3

Zinc 0.1

Molybdenum 0.2

FromAgroDynamicsPublications,Brooklyn,NewYork.

Table5.NutrientConcentrationforCucumbersinGrodanRockwool

pH 5.06.0

EC 2.0mmhos

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N 150ppm

P 35ppm

Ca 150ppm

Mg 30ppm

Fe 1.0ppm

Mn 0.75ppm

B 0.2ppm

Cu 0.2ppm

Zn 0.2ppm

Mo 0.03ppm

FromAgroDynamicsPublications,Brooklyn,NewYork.

Table6.DeficiencySymptomsfortheEssentialElementsinppm

Element Symptoms

Stuntedgrowth,foliagebecomesyellow(chlorotic)startinginolderleaves.Somecrops(corn,tomatoes)may
Nitrogen:
showareddishcolorinsteadofyellow.

Plantsaredwarfedwiththinstemsandsmallleaves.Anthocyaninpigmentsmayaccumulategivingplantsa
Phosphorus:
purplishcoloroccurringfirstinolderleaves.

Potassium: Olderleavesdevelopmarginalbrowningwhichcanextendintotheleaves,andforwardcurlingofleaves.

Shoottipsyellowanddieback,youngshootshaveabnormalgrowthwitheventualdieback.Newleaves
Calcium:
affectedfirstwithdistortedleafgrowthrootstipsdiebackleavingshortstubbyrootswithblackspots.

Magnesium: Interveinalchlorosisdevelopingfirstontheolderleaves.Witheringofoldleaves.

Sulfur: Yellowing(chlorosis)ofleavesusuallybeginninginnewleaves.Yellowingbecomesofftaninmanycrops.

Iron: Interveinalchlorosisbeginningonyoungerleaves.

Manganese: Interveinalchlorosisonleavesnearthetipoftheplant.Leavesmaydevelopnecroticlesionsanddrop.

Diebackofshootandroottips,stuntedgrowth.Internaltissuesmaydiscolororbecomehollowincauliflower
Boron:
andbeets.Leafsymptomsincludecurling,brittleness,wilting,chloroticspots.

Zinc: Shortenedinternodes,youngleavesaresmall,mayshowinterveinalchlorosis.

Molybdenum: Interveinalchlorosisbeginningonolderleavesmovinguptoyoungerleaves.

AdaptedfromResh,H.M.1983.HydroponicFoodProduction,2ndEdition.WoodbridgePressPublishingCo.,SantaBarbara,Ca.
pp.335andBergman,ErnestL.1985.NutrientSolutionCultureofPlants.ThePennsylvaniaStateUniv.CollegeofAgriculture,
ExtensionServiceHort.MimeoSeriesII:160.pp.21.

Table7.ToxicitySymptomsfortheEssentialElements

Nitrogen: Foliageisheavywithmanydarkgreenleaves,fewflowersandfruits.

Phosphorus: Rarelyoccurs,nosymptomsnoted.Mayleadtoadeficiencyofnitrogen,potassium,zincorcopper.

Potassium: Rarelyoccurs,expressedasmagnesiumdeficiency.

Sulfur: Reductioninleafsize,leavesmayshowinterveinalyellowing.

Calcium: Symptomsusuallyexpressedasdeficiencyofpotassium,magnesium,iron,manganese,orboron.

Magnesium: Poorgrowth.

Iron: Darkgreenfoliagemaydevelopmanganeseorzincdeficiency.

Manganese: Chloroticleaveswithunevenchlorophylldistributionanddarkbrownspots.

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Chlorine: Burningofleafmarginsbecomingnecroticintime,reducedleafsize.

Boron Youngerleavesdeformed,yellowingornecrosisofleavesbeginningatleaftip.

Zinc Mayresultinirondeficiency.

Copper Reducedgrowth,mayleadtoirondeficiency.

Molybdenum Rarelyoccurs,leavesoftomatoesturnyellow.

AdaptedfromResh,H.M.1983.HydroponicFoodProduction,2ndEdition.WoodbridgePressPublishingCo.,SantaBarbara,Ca.
pp.335.andBergman,ErnestL.1985.NutrientSolutionCultureofPlants.ThePennsylvaniaStateUniv.CollegeofAgriculture,
ExtensionServiceHort.MineoSeriesII:160.pp.21.

LiteratureCited

1.Arnon,D.I.andP.R.Stout.1939.Theessentialityofcertainelementsinminutequantityforplantswithspecialreferenceto
copper.PlantPhysio.14:371375.
2.Cooper,A.J.1973.Roottemperaturesandplantgrowth.IN:Res.Rev.4,CommonwealthBureauofHorticultureand
Plantationcrops.CommonwealthAgricultureBureau,England.
3.Bergman,E.L.1985.NutrientSolutionCultureofPlants.ThePennsylvaniaStateUniversityCollegeofAgriculture,
ExtensionServiceHort.MimeoSeriesII:160.pp.21.
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CourtesyoftheHydroponicSocietyofAmerica.Usedbypermission.

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