Ovule

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Thisarticleisabouttheplantstructure.Foranimalovules,seeOvum.
"Micropyle(botany)"and"Ovular"redirecthere.Forthestructureinanimals,seeMicropyle(zoology).Forthe
shape,seeoval.

LocationofovulesinsideaHelleborusfoetidusflower
Inseedplants,theovule("smallegg")isthestructurethatgivesrisetoandcontainsthefemalereproductivecells.
Itconsistsofthreeparts:Theintegument(s)formingitsouterlayer(s),thenucellus(orremnantof
themegasporangium),andfemalegametophyte(formedfromhaploidmegaspore)initscenter.Thefemale
gametophytespecificallytermedamegagametophyteisalsocalledtheembryosacinangiosperms.The
megagametophyteproducesaneggcell(orseveraleggcellsinsomegroups)forthepurposeoffertilization.After
fertilization,theovuledevelopsintoaseed.

Contents
[hide]

1Locationwithintheplant

2Ovulepartsanddevelopment
o

2.1Integuments,micropyleandchalaza

2.2Nucellus,megasporeandperisperm

2.3Megagametophyte

2.4Zygote,embryoandendosperm

2.5Typesofgametophytes

3Seealso

4References

5Bibliography

6Externallinks

Location within the plant[edit]

Infloweringplants,theovuleislocatedinsidetheportionoftheflowercalledthegynoecium.Theovaryofthe
gynoeciumproducesoneormoreovulesandultimatelybecomesthefruitwall.Ovulesareattachedtotheplacenta
intheovarythroughastalklikestructureknownasafuniculus(plural,funiculi).Differentpatternsofovule
attachment,orplacentation,canbefoundamongplantspecies,theseinclude:[1]

Apicalplacentation:Theplacentaisattheapex(top)oftheovary.Simpleorcompoundovary.

Axileplacentation:Theovaryisdividedintoradialsegments,withplacentasinseparatelocules.
Ventralsuturesofcarpelsmeetatthecentreoftheovary.Placentaearealongfusedmarginsofcarpels.Two
ormorecarpels.(e.g.Hibiscus,Citrus,Solanum)

Basalplacentation:Theplacentaisatthebase(bottom)oftheovaryonaprotrusionofthethalamus
(receptacle).Simpleorcompoundcarpel,unilocularovary.(e.g.Sonchus,Helianthus,Compositae)

Freecentralplacentation:Derivedfromaxileaspartitionsareabsorbed,leavingovulesatthecentral
axis.Compoundunilocularovary.(e.g.Stellaria,Dianthus)

Marginalplacentation:Simplesttype.Thereisonlyoneelongatedplacentaononesideoftheovary,as
ovulesareattachedatthefusionlineofthecarpel'smargins.Thisisconspicuousinlegumes.Simplecarpel,
unilocularovary.(e.g.Pisum)

Parietalplacentation:Placentaeoninnerovarywallwithinanonsectionedovary,correspondingto
fusedcarpelmargins.Twoormorecarpels,unilocularovary.(e.g.Brassica)

Superficial:Similartoaxile,butplacentaeareoninnersurfacesofmultilocularovary(e.g.Nymphaea)

Ovulestructure(anatropous)1:nucleus2:chalaza3:funiculus4:raphe
Ovuleorientationmaybeanatropous,suchthatwheninvertedthemicropylefacestheplacenta(thisisthemost
commonovuleorientationinfloweringplants),amphitropous,campylotropous,ororthotropous.
Ingymnospermssuchasconifers,ovulesareborneonthesurfaceofanovuliferous(ovulebearing)scale,usually
withinanovulatecone(alsocalledmegastrobilus).Insomeextinctplants(e.g.Pteridosperms),megasporangiaand
perhapsovuleswereborneonthesurfaceofleaves.Inotherextincttaxa,acupule(amodifiedleaforpartofa
leaf)surroundstheovule(e.g.CaytoniaorGlossopteris).

Ovule parts and development[edit]

Plantovules:Gymnospermovuleonleft,angiospermovule(insideovary)onright

Modelsofdifferentovules,BotanicalMuseumGreifswald
Theovuleappearstobeamegasporangiumwithintegumentssurroundingit.Ovulesareinitiallycomposed
ofdiploidmaternaltissue,whichincludesamegasporocyte(acellthatwillundergomeiosistoproduce
megaspores).Megasporesremaininsidetheovuleanddividebymitosistoproducethehaploidfemale
gametophyteormegagametophyte,whichalsoremainsinsidetheovule.Theremnantsofthemegasporangium
tissue(thenucellus)surroundthemegagametophyte.Megagametophytesproducearchegonia(lostinsomegroups
suchasfloweringplants),whichproduceeggcells.Afterfertilization,theovulecontainsadiploidzygoteandthen,
aftercelldivisionbegins,anembryoofthenextsporophytegeneration.Infloweringplants,asecondsperm
nucleusfuseswithothernucleiinthemegagametophyteformingatypicallypolyploid(often
triploid)endospermtissue,whichservesasnourishmentfortheyoungsporophyte.

Integuments,micropyleandchalaza[edit]
Anintegumentisaprotectivecelllayersurroundingtheovule.Gymnospermstypicallyhaveoneintegument
(unitegmic)whileangiospermstypicallyhavetwo(bitegmic).Theevolutionaryoriginoftheinnerintegument
(whichisintegraltotheformationofovulesfrommegasporangia)hasbeenproposedtobebyenclosureofa
megasporangiumbysterilebranches(telomes).[2]Elkinsia,apreovulatetaxon,hasalobedstructurefusedtothe
lowerthirdofthemegasporangium,withthelobesextendingupwardsinaringaroundthemegasporangium.This
might,throughfusionbetweenlobesandbetweenthestructureandthemegasporangium,haveproducedan
integument.[3]
Theoriginofthesecondorouterintegumenthasbeenanareaofactivecontentionforsometime.Thecupulesof
someextincttaxahavebeensuggestedastheoriginoftheouterintegument.Afewangiospermsproducevascular
tissueintheouterintegument,theorientationofwhichsuggeststhattheoutersurfaceismorphologicallyabaxial.
ThissuggeststhatcupulesofthekindproducedbytheCaytonialesorGlossopteridalesmayhaveevolvedintothe
outerintegumentofangiosperms.[4]
Theintegumentsdevelopintotheseedcoatwhentheovulematuresafterfertilization.
Theintegumentsdonotenclosethenucelluscompletelybutretainanopeningattheapexreferredtoas
themicropyle.Themicropyleopeningallowsthepollen(amalegametophyte)toentertheovuleforfertilization.

Ingymnosperms(e.g.,conifers),thepollenisdrawnintotheovuleonadropoffluidthatexudesoutofthe
micropyle,thesocalledpollinationdropmechanism.[3]Subsequently,themicropylecloses.Inangiosperms,onlya
pollentubeentersthemicropyle.Duringgermination,theseedling'sradicleemergesthroughthemicropyle.
Locatedoppositefromthemicropyleisthechalazawherethenucellusisjoinedtotheinteguments.Nutrientsfrom
theplanttravelthroughthephloemofthevascularsystemtothefuniculusandouterintegumentandfrom
thereapoplasticallyandsymplasticallythroughthechalazatothenucellusinsidetheovule.Inchalazogamous
plants,thepollentubesentertheovulethroughthechalazainsteadofthemicropyleopening.

Nucellus,megasporeandperisperm[edit]
Thenucellus(plural:nucelli)ispartoftheinnerstructureoftheovule,formingalayerofdiploid(sporophytic)
cellsimmediatelyinsidetheinteguments.Itisstructurallyandfunctionallyequivalenttothemegasporangium.In
immatureovules,thenucelluscontainsamegasporocyte(megasporemothercell),which
undergoessporogenesisviameiosis.Ingymnosperms,threeofthefourhaploidsporesproducedinmeiosis
typicallydegenerate,leavingonesurvivingmegasporeinsidethenucellus.Amongangiosperms,however,awide
rangeofvariationexistsinwhathappensnext.Thenumber(andposition)ofsurvivingmegaspores,thetotal
numberofcelldivisions,whethernuclearfusionsoccur,andthefinalnumber,positionandploidyofthecellsor
nucleiallvary.Acommonpatternofembryosacdevelopment(thePolygonumtypematurationpattern)includesa
singlefunctionalmegasporefollowedbythreeroundsofmitosis.Insomecases,however,twomegasporessurvive
(forexample,inAlliumandEndymion).Insomecasesallfourmegasporessurvive,forexamplein
theFritillariatypeofdevelopment(illustratedbyLiliuminthefigure)thereisnoseparationofthemegaspores
followingmeiosis,thenthenucleifusetoformatriploidnucleusandahaploidnucleus.Thesubsequent
arrangementofcellsissimilartothePolygonumpattern,buttheploidyofthenucleiisdifferent.[5]
Afterfertilization,thenucellusmaydevelopintotheperispermthatfeedstheembryo.Insomeplants,thediploid
tissueofthenucelluscangiverisetotheembryowithintheseedthroughamechanismofasexual
reproductioncallednucellarembryony.

Megagametophyte[edit]

MegagametophyteformationofthegeneraPolygonumandLilium.Triploidnucleiareshownasellipseswith
threewhitedots.Thefirstthreecolumnsshowthemeiosisofthemegaspore,followedby12mitoses.

Ovulewithmegagametophyte:eggcell(yellow),synergids(orange),centralcellwithtwopolarnuclei
(brightgreen),andantipodals(darkgreen)
Thehaploidmegasporeinsidethenucellusgivesrisetothefemalegametophyte,calledthemegagametophyte.
Ingymnosperms,themegagametophyteconsistsofaround2000nucleiandformsarchegonia,whichproduceegg
cellsforfertilization.
Infloweringplants,themegagametophyte(alsoreferredtoastheembryosac)ismuchsmallerandtypically
consistsofonlysevencellsandeightnuclei.Thistypeofmegagametophytedevelopsfromthemegasporethrough
threeroundsofmitoticdivisions.Thecellclosesttothemicropyleopeningoftheintegumentsdifferentiatesinto
theeggcell,withtwosynergidcellsbyitssidethatareinvolvedintheproductionofsignalsthatguidethepollen
tube.Threeantipodalcellsformontheopposite(chalazal)endoftheovuleandlaterdegenerate.The
largecentralcelloftheembryosaccontainstwopolarnuclei.

Zygote,embryoandendosperm[edit]
Thepollentubereleasestwospermnucleiintotheovule.Ingymnosperms,fertilizationoccurswithinthe
archegoniaproducedbythefemalegametophyte.Whileitispossiblethatseveraleggcellsarepresentand
fertilized,typicallyonlyonezygotewilldevelopintoamatureembryoastheresourceswithintheseedarelimited.
[citationneeded]

Infloweringplants,onespermnucleusfuseswiththeeggcelltoproduceazygote,theotherfuseswiththetwo
polarnucleiofthecentralcelltogiverisetothepolyploid(typicallytriploid)endosperm.Thisdoublefertilization
isuniquetofloweringplants,althoughinsomeothergroupsthesecondspermcelldoesfusewithanothercellin
themegagametophytetoproduceasecondembryo.Theplantstoresnutrientssuchasstarch,proteins,andoilsin
theendospermasafoodsourceforthedevelopingembryoandseedling,servingasimilarfunctiontotheyolkof
animaleggs.Theendospermisalsocalledthealbumenoftheseed.[citationneeded]
EmbryosmaybedescribedbyanumberoftermsincludingLinear(embryoshaveaxileplacentationandare
longerthanbroad),orrudimentary(embryosarebasalinwhichtheembryoistinyinrelationtotheendosperm).[6]

Typesofgametophytes[edit]
Megagametophytesoffloweringplantsmaybedescribedaccordingtothenumberofmegasporesdeveloping,as
eithermonosporic,bisporic,ortetrasporic.[citationneeded]

See also[edit]

Gynoecium

Ovum

Alternationofgenerations

Meiosis

Oogenesis

Placentation

References[edit]
1.

2.

Jumpup^Kotpal,Tyagi,Bendre,&Pande.ConceptsofBiologyXI.Rastogi
Publications,2nded.NewDelhi2007.ISBN8171338968.Fig.38Typesofplacentation,page2
127
Jumpup^Herr,J.M.Jr.,1995.Theoriginoftheovule.Am.J.Bot.82(4):54764

3.

^Jumpupto:

abStewart,W.N.;Rothwell,G.W.(1993).Paleobotanyandtheevolutionof
plants.CambridgeUniversityPress.ISBN0521382947.

4.

Jumpup^FrohlichandChase,2007.Afteradozenyearsofprogress,theoriginof
angiospermsisstillagreatmystery.Nature450:11841189(20December2007)
|doi:10.1038/nature06393;

5.

Jumpup^Gifford,E.M.;Foster,A.S.(1989),Morphologyandevolutionofvascular
plants,NewYork:W.H.FreemanandCompany

6.

Jumpup^TheSeedBiologyPlace:Structuralseedtypesbasedoncomparativeinternal
morphology

Bibliography[edit]

P.H. Raven, R.F. Evert, S.E. Eichhorn (2005): Biology of Plants, 7th Edition, W.H.
Freeman and Company Publishers, New York, ISBN 0-7167-1007-2

Peter K. Endress.Angiosperm ovules: diversity, development, evolution. Ann Bot (2011)

107 (9): 1465-1489. doi: 10.1093/aob/mcr120