Nutrientdeficiencycausesmorphologicalandphenotypicresponsesinwheatseedling

JulioFrancisco
FloridaInstituteofTechnology

Introduction:
Wheatisthemostproductiveandimportantcropwithover600milliontonnesof
wheatharvestedeveryyear,mainlyforhumanconsumptionandlivestockfeeding(1).
Furtherimprovementinwheatyieldisrequiretomeetthecurrentandimpending
challenges(2).Fosteringwheatplantcanbeaccomplishedbyprovidingadequate
amountofnutrientandwater.Inordertohaveabetterunderstandingoftheimportance
ofnutrients,onemustrecognizetheconsequencesofdeprivingsuchnutrient.The
followingreportprovidesabackgroundinformationontheroleofmacroand
micronutrients.Also,thisreportfurtherdiscussanexperimentconductedtowitnessthe
effectofdeprivingwheatofcertainnutrients.Lastly,thispaperaddressesthe
physiological,morphological,andphenotypicchangesforeachdeficiencies.
NitrogenFunctionandDeficiency
Nitrogenisamacronutrientthatsvitalinpromotingplantsgrowth,delaying
maturity,andenhancingthegreennessofplantsfoliage(3).Nottomention,nitrogen
greatlyinfluencethedryweightsoftheshootandrootofaplant,leafstomatal
conductance,transpirationrate,photosyntheticrate,andabsorbancerateof
micronutrients(e.g.zinc,manganeseandcopper)(30).
Evidencedemonstratesthatsomesymptomsofnitrogendeficiencyinwheat
includesdelayingrowthandchlorosisinwheatplant(4).Furtherstudiesalsoindicated
thattherateofsenescenceincreasedduringnitrogendeficiency(5).Nottomention,the
rateofrootelongationdecreasedwhichenabletoaccessnitrogenresourcefromthe

ground.Withthisbeingsaid,onecanhypothesisthatstemheightwoulddecreasesand
thefibrousrootlengthdecreases.Also,onewouldobservechlorosisdisorderand
witheringoftheleave.
PotassiumFunctionandDeficiency
Potassiumactascofactorinmanyenzyme,andusedestablishingturgor
pressureandmaintaincellelectroneutrality(3).
Ingeneral,plantthatendurepotassiumdeficiencyshowedsignsofmarginal
chlorosiswhichdevelopintonecrosisatthetipoftheleave(3).Furthermore,studies
areinconsistentwiththeeffectofPotassiumonrootdevelopment.Ashley,Grant,and
Grabov(2005)showedthatPotassiumdeficiencyZeamaysunderwentmorphological
changessuchastriggeringrootdevelopment(31).Ontheotherhand,Drew(1975)
demonstratedthatbarleyplantdevelopedpoorlateralrootduringpotassiumdeprivation
(14).Therefore,nohypothesiscanbesynthesisabouttheeffectofdeprivingPotassium
onplant.Onecanhypothesisthatthestemheightwoulddecreaseduetothedecrease
inboththestomatalconductanceandphotosyntheticratewhichresultinginlimiting
growth.
PhosphorusFunctionandDeficiency
PhosphorusisessentialtoprovideenergyintheformofATPtodrive
biochemicalprocesssuchasphotosynthesis(33)andistheoneofthemajor
componentsingeneticinformation(DNAandRNA).Itisnotobviousifaplantis
experiencingphosphorusdeficiencyorsomeothercondition.Therefore,diagnosinga
plantbasedongeneralappearanceisdifficult.Whenthereisalowlevelofphosphorus,

theplantusuallyexperiencemorphologicalchangesuchasanincreasedintheroot
surface,whichenabletheplanttogainbetteraccesstophosphorusinthesoil.Notto
mention,primaryrootisreducedandthelengthanddensityofroothairincreases.
Furthermore,theelongationrateofaxileanddensityofrootceasetochangeduring
phosphorusdeprivationinmaizehowever,newaxilerootemerges(34).Furthermore,
phosphorusdeficientplantischaracterizedbytheirdarkgreencolorationwith
reddishpurpleleaftipsandmargin(34).Inaddition,theleafarea,leafelongationrate,
andgrainyieldarereducedalongwithdelayinleafandfloweringdevelopment(33).
Overall,onecanhypothesisthatthenumberandsizeofrootwouldincrease.
SulfurFunctionandDeficiency
Sulfurisanessentialelementtosynthesisamino,protein,vitamins,and
coenzymes(35).Inaddition,sulfurisrequireforferredoxinstosynthesizechlorophyll
(35).Usually,thereisanincreaseinyield,size,andweightwhenagrainareexposedto
excessiveamountofsulfur.General,sulfurdeficiencyischaracterizedbychlorosis,
stuntgrowth,andanthocyaninaccumulation(3).Symptomsofsulfurdeficiency
includinginterveinalchlorosisandyellowingleaves,appearinyoungcornplant(3).
Moreover,sulfurdeficiencyinducesgeneticchangescauseanimprovementinsulfate
uptake,reductioncapacity,enhancementofsulphateremobilizationfromthevacuole
(36).
CalciumFunctionandDeficiency
Calciumisimportantforcellwallelasticityandexpansion,mitoticspindleduring
celldivision,andimplicatedasasecondmessengerforplantresponse(3).Itis

responsibleforgrowthatthetipoftheplant.Nottomention,calciumassistinthe
developmentofroothairandstarchtransport(37).Usually,youngplantwithcalcium
deficiencyexhibitdeformedleaves.Inaddition,plantgrowshortandstubbyrootswhen
deprivedofcalcium.
MagnesiumFunctionandDeficiency
Magnesiumplaysarolephosphatetransferandconstitutechlorophyllmolecule.When
aplantismagnesiumdeficient,theveinsbecomedarkgreenwithyellowareasinthe
middleoftheveininaphenomenoncalledinterveinalchlorosis.Symptomalsoinclude
prematureleafabscission.Therefore,onecanhypothesisethatthestemheightwould
bereducedandshowchlorosis.
Limitedamountofresearcheswereconductedontheimpactofdeprivingwheat
ofmacroandmicronutrient.Withthisinmind,afirsthandapproachwasundertakento
determinethecharacterizationofthestemheight,fibrousrootlength,adventitiousroots,
andweightwhenstarvingwheatplantofessentialnutrients.Overall,onecan
hypothesisthateachnutrientwillcausesvariousofuniquechangesonthe
morphologicalandphenotypicstructureofthewheatplant.
ExperimentalProcedures:
Thisstudywasperformedina6weekstimespaninwhichwheatseedlingswere
deprivedofcertainnutrients.Thewheatplantreceivednutrientvianutrientenhanced
water.Thefollowingtreatmentswereestablished:apositivecontrol(receivedall
essentialnutrients),nitrogendeficiency,potassiumdeficiency,phosphorusdeficiency,
sulfurdeficiency,calciumdeficiency,magnesiumdeficiency,and

micronutrientdeficiency.Priortoperformingtheexperiment,theseedswereallowedto
pregerminatefortwodaysinwhichthepregerminatedseedswateredandplacedin
thedark.Next,threetofivepregerminatedwheatseedsweregrowninthesamepotfor
eachtreatment.Importanttomention,thesoilpotwas1:1:1ratioconsistingofsand,
perlite,andvermiculite.Table1demonstratetheconcentrationofnutrientgivento
individualplants.
Table1.Thetypeofnutrientdeficiencywiththecorrespondingamountof
chemicalcompoundsdilutedin1000mLsolution.
Solution

10mL

10mL

10mL

10mL

2mL

10mL

Control

Ca(NO3)2

KNO3

KH2PO4

MgSO4

FeEDTA

0.5MMicronutrient

Nitrogen

CaCl2

KCl

KH2PO4

MgSO4

FeEDTA

0.5MMicronutrient

Potassium

Ca(NO3)2

(5mL)KNO3

NaH2PO4

MgSO4

FeEDTA

0.5MMicronutrient

Phosphorus

Ca(NO3)2

KNO3

(2mL)KCl

MgSO4

FeEDTA

0.5MMicronutrient

Sulfur

Ca(NO3)2

KNO3

KH2PO4

MgSO4

FeEDTA

0.5MMicronutrient

Calcium

NaNO3

KNO3

KH2PO4

MgSO4

FeEDTA

0.5MMicronutrient

Magnesium

Ca(NO3)2

KNO3

KH2PO4

FeEDTA

0.5MMicronutrient

Micronutrient

Ca(NO3)2

KNO3

KH2PO4

MgSO4

FeEDTA

Furthermore,eachplantweregrowninasemihydroponicenvironmentand
placedunderthesamegreenhouseconditions(e.g.samehumidity,temperature,Etc).
Afterthe6weeks,thestemheight,rootlength,numberofroot,andgeneralappearance
oftheplantweredetermined.Theheightandlengthwasmeasuredusingaruler,while
theweightwasmeasuredwithabalancingscale.
Eachdeficiencytypewascomparedtothenormal(control)growingwheatplantto
determineiftheplantincreasedordecreasedinmeanstemheight,rootlength,and

numberofrootsrelativetothecontrol.Also,thephysiologicalandmorphological
mechanismoccurredduringnutrientdeprivationwillbediscuss.
Results:
Inthecontrolpot,nodefectwasdetected(Fig.1).

(a)

(b)

Figure1.Control.Theentireplant(a&b)after6weeks.

However,thenitrogendeficiencywheatplantshoweddelayedinmaturitywhen
comparingtoothernutrientdeficientindividuals(Fig.2).Inaddition,Theentireleaf
havedevelopedchlorosis(abnormallyyellowcolorinplanttissue)andexhibita
reductioninstemheight(incomparisonwiththecontrolplants).

(a)(b)(c)

Figure2.NitrogenDeficient.Thefollowingfiguresshowtheentireplant(a)alongwitha
closeupofthestem(b&c)after6weeksofbeingdeprivedofnitrogen.
Furthermore,thepotassiumdeficientwheatplantexhibitedayellowbrownnecrosisat
thetipoftheleavesalongwithmarginalchlorosis(Fig.3).Inaddition,theplantdidnot
producedprimaryroots,butgeneratedmanysimilarsizelateralroots.

(a)
(b)
Figure3.PotassiumDeficient.Thestem(a)androot(b)ofthepotassiumdeficientwheat
plantafter6weeks.

Thephosphorusdeficientwheatshowedchlorosisdefectatthetipoftheleaves,whilethe
restoftheplantdidnotexhibitanyfurtherchange(Fig.4).

(a)
(b)
Figure4.PhosphorusDeficient.Thephenotypicalchangeswhichoccurredonthestem(a)
androot(b)ofwheatafter6weekswithoutphosphorus.
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Thesulfurdeficientplantexhibitstuntstemgrowthandincreasedinrootelongation(Fig.5).
Nottomention,someleavesoftheplantexperiencedsenescence(nolongerabletodivide
butmetabolicallyactive)fromthetipoftheleafnearthebaseoftheplant.

(a)
(b)
Figure5.Sulfurdeficiency.Thestem(a)andtheroot(b)ofthesulfurdeficientwheatplant.

Deprivingcalciumcausedtheplanttoexhibitchlorosisonthemarginandtipofthestem(Fig.
6).Suchabnormityappearedintheearlystageofdevelopment.Inaddition,thestemsofthe
plantwerepoorlydeveloped,whichcausedthestemtotiltatanangleinsteadofgrowing
vertically.

(a)
(b)
(c)
Figure6.CalciumDeficiency.Thephenotypicandmorphologicalchangestothestem(a),
node(b),androot(c)ofwheatplantafterweek6withoutcalcium.
9


Inthemagnesiumdeficiencywheat,therewereabout5seedsgrowninthepotandonlyone
individualplantdeveloped(Fig.7).Inaddition,thetipandmarginofthestemexhibitchlorosis.

(a)
(b)
Figure7.MagnesiumDeficient.Theentireplant(a)andtheroot(b)ofwheatplantafterweek
6.

Inthecalciumdeficientwheat,novisibledefectwasdetectedduringtheplantgrowthand
development(Fig.8).

(a)
(b)
Figure8.MicronutrientDeficient.Theentireplant(a)andtherootofthe
micronutrientdeficientwheatafterweek6.

10

Afterthe6weekstimespan,thenitrogendeficientplantpossessedthelowest
meanstemheightamongalltreatments,whilethepotassiumdeficientthegreateststem
height(Fig.10).Furthermore,thepotassiumpossessedtheshortestfibrousroot,while
themagnesiumproducedthelongestfibrousroot.Lastly,micronutrientpossessedthe
highestweight,whilenitrogenpossessedthelowestweight.Overall,theKdeficiency
wheatplantendurethegreatestmorphologicalchangesinthisexperiment.

11

Figure10.Themeanstemheight(incm),fibrousrootlength,andweightforeachtypeof
deficienciesafter6week.

Discussion:
Thepurposeofthisinvestigationwastodetermineifcertainnutrientdeficiencyinduces
uniquemorphologicalandphenotypicalchangestothestemandrootsystem.The
followingdiscussiongivenanoveralloftheresultoftheinvestigationandprovidefurther
evidencetothecauseofsuchchanges.
EffectofNitrogenDeficiency
Ndeficientplantdemonstratedsignofchlorosisandexhibitedsenescence.
Evidencesconfirmthatdevelopedwheatplantwouldsufferfromsuchdiscoloration
(4,5).Thisisduetothebreakdowninchlorophyllproduction,causingchlorosis
symptomstofirstmergefromthetip,thenextendingdownwardtotheleaf(5).Further
studiesindicatedthatNdeficiencytriggersareductionRuBPregenerationandincrease
theactivationofRuBPCOactivity,whichoverallrestrictphotosynthesisandfurther
contributetochlorosisintheleaves(6).Inaddition,itwasreportedthatyoungwinter
12

wheatplantcanalsoexhibitpurplecoloredleavesduetoexpressionofgenecontrolling
coleoptiles(4).
Furthermore,theresultwasconsistentwithpreviousstudiesinwhichNdeficient
increasedtherateofleafsenescence(5).Studiesindicatedthatsenescenceprocess
mainlyoccurredwhentheleafreachesitsmaximumsize,inwhichthereisadeclinein
thephotosynthesisassociatedproteinanddegradationofchloroplastenzyme(7).
Moreover,Ndeficientplantdemonstratedreductioninfibrousroot.Thisresultis
notconsistentwithotherstudies(8,9).Teplova,Veselov,andKudoyarova(1998)
demonstratedthatNdeficiencypromotedrootelongationandshootretardinwheat
plant(8).Onecanpostulatethattherootelongationenableplanttogainbetteraccess
tonitrogeninthesoilduringNdeprivation.
EffectofPotassiumDeficiency
Potassiumplaysavitalroleinrateofstomatalconductance,inwhichK+fluxinto
andoutofthestomatalguardcells.ThisfluctuationofK+affectstheosmoticpotential
withinthecell,whichdetermineifthestomatalguardcellsopenorclose.During
potassiumdeficiency,thestomatalconductanceratedecreases,whichdecreases
photosyntheticprocess(10).Thisdecrementintherateofphotosynthesiscanbeseen
bythechlorosisinthestemofwheatplant(11,5).Inthisexperiment,Kdeficientleaves
exhibitedchlorosisatthelateralandinterveinalsideofthestem.
Moreover,thestemheightwasslightlyabovethecontrolgroup.Thisisnot
consistentwiththeresultofotherstudieswithotherspeciesofplant.Forinstance,
Masakaa,Mugutia,&Chivandib(2008)underlinedthatKdeficienttobaccoplants

13

showed86%reductionincomparisontothecontrol,whichreceivedacompletenutrient
solution(12).
Inaddition,Kdeficientplantcontainedmorelateralrootthanprimaryroots.Not
tomention,theKdeficientplantrootspossesstheshortestfibrousrootthananyother
nutrientdeficientplant.Studiesshowedsimilarmorphologicalresponsewithbarleyand
Arabidopsis(13,14).DuringKdeprivation,acascadeofcellularactivitiesoccur.First,
thereisanincreasedofgenesforethyleneproduction,inwhichethyleneinhibitsroot
growth.Secondly,Hak5,AtKC1,andKEA5genesareupregulatedtoencodeforK+
transportproteinsinordertoenhanceK+ionuptakeintotherootcell.Lastly,reactive
oxygenspecies(ROS),suchasH2O2,increasesincertainregionoftheplant,inwhich
ROSactivelyperformedK+uptakeandtranslocation(15).Overall,theplantwould
exhibitreductioninrootsizeandundergogeneticandphysiologicalchangesduring
Kdeprivation.
EffectofPhosphorusDeficiency
Incomparisontothecontrol,Pdeficientplantexperiencedreductionofstem
heightandweight,yetthefibrousrootlengthincreased.Resultswereconsistentwith
report(5).ReductioninstemheightcanbeexplainedbythefactthatPdeficiencylimits
celldivisionattheshootapicalmeristemand,thus,inhibitingnewlydevelopedleaves
(16).
Inaddition,Sarker,Karmoker,andRashid(2010)determinedthatthesizeand
numberofstomatawerereducedinZ.maysalongwithdecreasedsizeofguardcells,
leadingtoadecreaseinZ.maysgrowth(17).

14

Furthermore,phosphorusdeficientwheatplantexhibitedchlorosisdisorder.This
maybealsoduetoreductionofstomatalconductancerate,leadingtodecreaseinCO2
uptakeand,thus,restrictingmetabolismfortheplant.Also,Plenet,Etchebest,Mollier,
andPellerin(2000)demonstratedthatleafelongationrateisreducedduringP
deprivation(18).
Inaddition,theincreasedlengthofthefibrousrootenableplanttoincrease
surfaceareainordertoincreaseabsorbancerate(19).MollierandPellerin(1999)
determinethatthemaizerootwereslightenhanced,butreducedaftercertainperiodof
time(19).
EffectofSulfurDeficiency
Inthisinvestigation,deficiencyofsulfurinducedaccelerativesenescenceofthe
leavesandincreaserootlengthalongwithstuntgrowth.Sdeficiencytriggersa
reductioninRubiscoactivityand,thus,decreaseinchlorophyllcontent.Asaresult,
chlorosisappearsinyoungleaves(20).
Furthermore,thefibrousrootlengthandweightoftheplantincreased.Thisis
duetothefactthatgenesencodingforauxinproductionisoverexpressedand
accumulatesduringsulfurdeficiency(21).Rootgrowthinducedbyanauxicalledindole
aceticacid(IAA)alongwithpositiveregulationofauxinresponsivegenesasseenin
Arabidopsisspecies(22).
EffectofCalciumDeficiency
Calciumdeficientwheatplantexhibitedlightyellowcolorationatthetipand
marginoftheleaves.Suchabnormalitieswasalsoseenduringwheatearly

15

development.Itisreportthatonlyyoung,newlygrownwheatexemplifychlorosis
characteristic,whilemature,developedwheatretaintheirdarkgreencolorduringCa
deficiency(5)
Inaddition,thestemheightandfibrousrootlengthwereroughlysimilartothe
controltreatment.Thisresultisnotconsistentwithotherstudies.LascarisandDeacon
(1991)statedthatcalciumdeficiencyenhancetherateofrootcorticaldeath(RCD),
causingtherootandshoottobecomereduce(23).
EffectofMagnesiumDeficiency
Whenwheatplantenduredmagnesiumdeficiency,therewasareductioninseed
germinationandstemheight.However,thefibrousrootlengthandweightincreased.
Also,thewheatplantexhibitchlorosisatthetipoftheleaf.
Furthermore,chlorosisandreducedshootsystemisconsistentwithotherstudies
(24,5).Thecauseofchlorosisisbythedisruptionofthephotosyntheticprocess,
particular,bythedecrementoftheHillreactionactivityandRuBPcarboxylaseactivity
(25)
Studiesdonotsupporttheresultoftheexperiment,inwhichthefibrousroot
lengthandweightincreased.IthasbeenreportthatMgdeficientwheatplantendure
restrictedrootgrowth(24,5).
EffectofMicronutrient
Themicronutrientdeficientwheatdidnotdemonstrateanyvisiblesignsor
symptomsinthestemorrootoftheplant.Inaddition,thestemheightwassimilartothe
controltreatment.However,thelengthofthefibrousrootandweightoftheplant

16

increasedincomparisonthecontrolgroup.Itisdifficulttodistinguishwhich
micronutrient(e.g.Fe,Cl,Cu,CO,Mg,Mn,B,Mo,andZn)wouldhaveanadverse
effectonthegrowthanddevelopmentofwheatplant.Manyliteraturestudiesfocuson
theeffectoneparticularmicronutrientdeprivation.Inaddition,limitamountofstudies
wereperformedondeprivingplantofallmicronutrients.Eachnutrientdeficiencieshave
variouseffectonthegerminationofwheat.Somemineraldeficiencyhavecontradicting
effects.Forexample,lowboronconcentrationhasbeenshowtoincreaseweightof
shootandrootsystemofwheatplant(26).Onthecontrary,Kleli,Eker,andCakmak
(2004)showedthatzndeficiencydecreaseshootweightinwheat(27).Therefore,itis
challengingtodeterminewhichmineralshaveagreaterinfluenceonthegrowthand
developmentofwheatplant.
Furtherimprovementcanbemadeforfutureexperiment.First,theplantswere
notgiventhesameamountofnutrientenhancedwater.Givingaplantmoreorless
nutrientfilledwatercouldrestrictplantgrowthanddevelopment(3).Nottomention,
LawsonandBlatt(2014)statedthatthestomataofwellwateredplants...reduce
photosyntheticrates,whichfurtherreducestheplantabilitytogrow(28).Forthese
reason,adiscreteamountofwatermustbeestablish.Secondly,alargersamplesizeis
requiretodetermineiftheexperimentisvalid.Inthisexperiment,onlyonesamplepot
wasusedforeachdeficiencies(includingthecontrol).Thisdoesnotgiveaexcellent
representationofallwheatplantand,therefore,additionalsamplemustbeincluded.
Thirdly,thewheatplantwerenotfullydevelopedduetotimeconstraint.Inorderto
possessanaccurateexamination,thewheatplantshouldbeallowtodevelopintoa

17

matureplant.Nottomention,somedeficiencies,suchaspotassium,donotappearuntil
maturity(29).Therefore,thetimespanoftheexperimentshouldincrease.
Allandall,nutrientdeficiencycaninduceuniquephysiologicalandmorphological
changes.Suchchangescanbeseenvialeafappearancealongwithmorphologicaland
phenotypicalchangesoftherootandshootsystem.Futureinvestigationshouldinclude
additionalcontrol,alargersamplesize,andanextensiononthedurationtoallow
maturity.Inaddition,furtherexperimentshouldbeconductedonthedifferent
concentrationforeachmacroandmicronutrient.
Lastly,studyofnutrientdeficientwheatplantprovideanopportunitytoimprove
agriculturalmethodsandincreasecropproductivity.Forinstance,thereisachallenge
foragriculturetomeettheworldsincreasingdemand.Tomeetsuchchallengerequire
touptakemorelandforfurthercultivationortoincreaseyieldunderrestrictedarea.
Impropermanagementofplantnutrienthavefurtherprovoketheproblem.Therefore,
understandingtheconsequenceofnutrientdeficiencycanprovokepeopletomakea
change.

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