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Fertilization
Embryology5Apr2016
title=Fertilization)

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Contents
1 Introduction
2 SomeRecentFindings
3 Objectives
4 Movies
5 FertilizationPreparation
6 Oogenesis
7 MenstrualCycle
8 Ovulation(HPGAxis)
9 ZonaPellucida
10 CoronaRadiata
11 GameteformationSpermatogenesis
11.1 Spermiogenesis
12 Ejaculate
13 FertilityWindow
14 FertilizationSite
14.1 FertilizationSpermatozoa
14.2 FertilizationOocyte
15 SpermatozoaOocyteInteraction
15.1 MembraneFusion
15.2 Izumo
15.3 Juno
15.4 CD9
15.5 Ovastacin
16 FormationoftheZygote
16.1 Mitochondria
17 SexDetermination
18 FertilizationProteinChanges
18.1 ProteinExpressionClassifiedbyMolecularFunctions
19 Abnormalities
19.1 CompleteHydatidiformMole
19.2 PartialHydatidiformMole
19.3 Parthenogenesis
19.4 Trisomy21
20 References
20.1 Textbooks
20.2 Reviews
20.3 Articles
20.4 SearchPubmed
21 AdditionalImages
22 ExternalLinks
23 GlossaryLinks

Introduction
Fertilizationisthefusionofhaploidgametes,eggandsperm,toformthediploidzygote.Notethoughtherecan
besubtledifferencesinthefertilizationprocesswhichoccursnaturallywithinthebodyorthroughreproductive
technologiesoutsidethebody,theoverallproductinbothcasesisadiplodzygote.Infertilizationresearch,
afterhumansthemouseisthemoststudiedspeciesfollowedbydomesticandfarmanimals.Theprocessof
fertilizationinvolvescomponentsof,andsignalingbetween,bothsperm(spermatozoa)andegg(oocyte).
Inadditiontoinvivofertilizationtherearemanynewinvitrotechnologiesrelatedtohumaninfertility(Assisted
ReproductiveTechnology(/embryology/index.php/Assisted_Reproductive_Technology))andanimalproduction
somaticcellnucleartransfer(SCNT)togenerateazygote.
NotedifferentspellingUSAspelling"Fertilization",Australianspelling"Fertilisation".

(/embryology/index.php/File:Human_fertilization_movie_2_fram
HumanFertilization
FertilizationLinks:Fertilization|Oocyte(/embryology/index.php/Oocyte_Development)|Spermatozoa
(/embryology/index.php/Spermatozoa_Development)|CellDivisionMeiosis
(/embryology/index.php/Cell_Division__Meiosis)|Zonapellucida(/embryology/index.php/Zona_pellucida)
|Zygote(/embryology/index.php/Zygote)|LectureFertilization(/embryology/index.php/Lecture__Fertilization)|CellDivisionMitosis
(/embryology/index.php/Cell_Division__Mitosis)|LectureWeek1and2(/embryology/index.php/Lecture__Week_1_and_2_Development)|Hydatidiform

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Mole(/embryology/index.php/Abnormal_Development__Hydatidiform_Mole)|AssistedReproductive
Technology(/embryology/index.php/Assisted_Reproductive_Technology)|LectureGenitalDevelopment
(/embryology/index.php/Lecture__Genital_Development)|MenstrualCycle
(/embryology/index.php/Menstrual_Cycle)|Morula(/embryology/index.php/Morula)|Blastocyst
(/embryology/index.php/Blastocyst)

(/embryology/index.php/File:Humanspermatozoa_EM01.jpg)

SomeRecentFindings
Versatileactionofpicomolargradientsofprogesteroneondifferentspermsubpopulations[1]"High
stepconcentrationsofprogesteronemaystimulatevariousspermphysiologicalprocesses,suchas
primingandtheacrosomereaction.However,approachingtheegg,spermatozoafaceincreasing
concentrationsofthehormone,asitissecretedbythecumuluscellsandthenpassivelydiffusesalongthe
cumulusmatrixandbeyond....Theresultssuggestaversatileroleofthegradualdistributionofverylow
dosesofprogesterone,whichselectivelystimulatetheprimingandtheacrosomereactionindifferent
spermsubpopulations."
JunoistheeggIzumoreceptorandisessentialformammalianfertilization"Fertilizationoccurs
whenspermandeggrecognizeeachotherandfusetoformanew,geneticallydistinctorganism.The
molecularbasisofspermeggrecognitionisunknown,butislikelytorequireinteractionsbetweenreceptor
proteinsdisplayedontheirsurface.Izumo1isanessentialspermcellsurfaceprotein,butitsreceptoron
theegghasnotbeendescribed.Hereweidentifyfolatereceptor4(Folr4)asthereceptorforIzumo1on
themouseegg,andproposetorenameitJuno.WeshowthattheIzumo1Junointeractionisconserved
withinseveralmammalianspecies,includinghumans.FemalemicelackingJunoareinfertileandJuno
deficienteggsdonotfusewithnormalsperm.RapidsheddingofJunofromtheoolemmaafterfertilization
suggestsamechanismforthemembraneblocktopolyspermy,ensuringeggsnormallyfusewithjusta
singlesperm."Nature16April2014
(http://www.nature.com/nature/journal/vaop/ncurrent/full/nature13203.html)
Nongeneticcontributionsofthespermnucleustoembryonicdevelopment[2]"Recentdatafrom
severallaboratorieshaveprovidedevidencethatthenewlyfertilizedoocyteinheritsepigeneticsignals
fromthespermchromatinthatarerequiredforproperembryonicdevelopment.Forthepurposesofthis
review,thetermepigeneticisusedtodescribealltypesofmolecularinformationthataretransmittedfrom
thespermcelltotheembryo.Thereareatleastsixdifferentformsofepigeneticinformationthathave
alreadybeenestablishedasbeingrequiredforproperembryogenesisinmammalsorforwhichthereis
evidencethatitmaydoso.Theseare(i)DNAmethylation(ii)spermspecifichistones,(iii)other
chromatinassociatedproteins(iv)theperinuclearthecaproteins(v)spermbornRNAsand,thefocusof
thisreviewand(vi)theDNAloopdomainorganizationbythespermnuclearmatrix.Theseepigenetic
signalsshouldbeconsideredwhendesigningprotocolsforthemanipulationandcryopreservationof
spermatozoaforassistedreproductivetechnologyasnecessarycomponentsforeffectivefertilizationand
subsequentembryodevelopment."
CD9tetraspaningeneratesfusioncompetentsitesontheeggmembraneformammalian
fertilization [3]"CD9tetraspaninistheonlyeggmembraneproteinknowntobeessentialforfertilization.
Toinvestigateitsrole,wehavemeasured,onauniqueacrosomereactedspermbroughtincontactwith
anegg,theadhesionprobabilityandstrengthwithasensitivityofasinglemoleculeattachment.Probing
thebindingeventsatdifferentlocationsofwildtypeeggwedescribeddifferentmodesofinteraction.Here,
weshowthatmoregameteadhesioneventsoccuronCd9nulleggsbutthatthestrongestinteraction
modedisappears.WeproposethatspermeggfusionisadirectconsequenceofCD9controlledsperm
eggadhesionproperties.CD9generatesadhesionsitesresponsibleforthestrongestoftheobserved
gameteinteraction.Thesestrongadhesionsitesimpose,duringthewholeinteractionlifetime,atight
proximityofthegametemembranes,whichisarequirementforfusiontotakeplace.TheCD9induced
adhesionsiteswouldbetheactuallocationwherefusionoccurs."
Gameterecognitioninmicedependsonthecleavagestatusofanegg'szonapellucida
protein [4]"spermeggrecognitiondependsonthecleavagestatusofZP2andthatbindingatthesurfaceof
thezonaisnotsufficienttoinducespermacrosomeexocytosis."

(/embryology/index.php/File:Early_zygote_labelled.jpg)
EarlyHumanZygote

(/embryology/index.php/File:Mouse
fertilization_01.jpg)
Thefirstpolarbodydeformsthemammalian
eggawayfromitsencapsulatingzona
pellucida

Morerecentpapers [Expand]

Objectives
Understandthemechanismsofgameteformation.
Understandthemechanismsofcelldivision.
Describethedifferencesbetweenmitosisandmeiosis.
Understandthemechanismsoffertilization,bothinvivoandinvitro.
Describethecleavageofthezygote.
HaveapreliminaryunderstandingoftheroleandprocessinmalesexdeterminationandXinactivation.
Understandtheabnormalitiesthatoccurduringthisperiodofdevelopment.

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Movies

(/embryology/index.php/Ovulation_Movie)
Ovulation
Page(/embryology/index.php/Ovulation_Movie)
|Play
(/embryology/images/0/00/Follicle_001.mp4)

(/embryology/index.php/Spermatozoa_Structure_Movie) (/embryology/index.php/Human_Spermatozoa_Motility_Movie)(/embryol
Spermatozoa
SpermatozoaMotility

Page
Page
Page
(/embryology/index.php/Spermatozoa_Structure_Movie) (/embryology/index.php/Human_Spermatozoa_Motility_Movie)(/embryol
|Play
|Play
Play
(/embryology/images/5/5b/Spermatozoa_animation.mp4)(/embryology/images/a/ac/Spermatozoa_motility_01.mp4)
(/embryol

(/embryology/index.php/Fertilization_Movie)
F ertilization
(/embryology/index.php/Mouse_Fertilization_Movie)
Page
MouseFertilisation
(/embryology/index.php/Fertilization_Movie)|
Page
Play
(/embryology/index.php/Mouse_Fertilization_Movie)|
(/embryology/images/5/5f/Fertilization_003.mp4)Play(/embryology/images/c/c0/Fertilization_001.mp4)

(/embryology/index.php/Pronuclear_Fusion_Movie)
PronuclearFusion
Page(/embryology/index.php/Pronuclear_Fusion_Movie)|
Play(/embryology/images/9/95/Pronuclear_fusion_001.mp4)

(/embryol

Page
(/embryol

FertilizationPreparation
Priortothefertilizationprocesscommencingboththegametesoocyte(egg)andspermatozoa(sperm)requirecompletionofanumberofbiologicalprocesses.
OocyteMeiosiscompletesMeiosis1andcommencesMeiosis2(arrestsatMetaphaseII).
SpermatozoaCapacitationfollowingrelease(ejaculation)andmixingwithotherglandularsecretions,activatesmotilityandacrosomepreparation.
MigrationbothOocyteandSpermatozoa.
oocyteovulationandreleasewithassociatedcells,fromovaryintofimbriathenintouterinetube(oviduct,uterinehorn,fallopiantube)andepithelialcilia
mediatedmovement.
spermatozoaejaculation,depositedinvagina,movementoftailto"swim"inuterinesecretionsthroughcervix,uterinebodyandintouterinetube,have
approximately2472htofertilizeoocyte.
EndocrinologyDiagramofthecomparativeanatomyofthemaleandfemalereproductivetracts(http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?
book=endocrin&part=A972&rendertype=box&id=A1230)

Oogenesis
Processofoogoniamatureintooocytes(ova,ovum,egg)
alloogoniaformprimaryoocytesbeforebirth,thereforeamaturationofpreexistingcellsinthefemale
gonad,ovary

(/embryology/index.php/File:Human_ovary_non

growing_follicle_model.jpg)
humansusuallyonly1ovumreleasedeverymenstrualcycle(IVFsuperovulation)
oocyteanditssurroundingcells=follicle
primary>secondary>ovulationreleases
OvaryHistologywholetransversesection(cortex,medulla)

(/embryology/index.php/File:Ovary5x.gif)
HistologyoftheOvary

MenstrualCycle
PrimaryOocytearrestedatearlyMeiosis1
diploid:22chromosomepairs+1pairXchromosomes(46,XX)
autosomesandsexchromosome
Oogenesispreantralthenantralfollicle(Graafianfollicleismatureantralfolliclereleased)
Secondaryoocyte
1Daybeforeovulationcompletes(stimbyLH)Meiosis1
haploid:22chromosomes+1Xchromosome(23,X)
nondisjunctionabnormalchromosomesegregation
beginsMeiosis2andarrestsatmetaphase
notenointerphasereplicationofDNA,onlyfertilizationwillcompleteMeiosis2

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Ovulation(HPGAxis)

(/embryology/index.php/File:Ova41he.jpg)
PreantralFollicle

(/embryology/index.php/File:Menstrual_cycle.png)
Hypothalmusreleasesgonadotropinreleasinghormone(GRH,luteinizinghormonereleasinghormone,
LHRH)>Pituitaryreleasesfolliclestimulatinghormone(FSH)andlutenizinghormone(LH)>ovaryfollicle
developmentandovulation.
releaseofthesecondaryoocyteandformationofcorpusluteum
secondaryoocyteencasedinzonapellucidaandcoronaradiata
Ovulationassociatedwithfollicleruptureandampullamovement.

ZonaPellucida
glycoproteinshellZP1,ZP2,ZP3,ZP4(mouseonlyZP13)
mechanicalprotectionofegg
involvedinthefertilizationprocess
spermbinding
adhesionofspermtoegg
acrosomereaction
releasesenzymestolocallybreakdown
blockofpolyspermy
alteredtopreventmorethan1spermpenetrating
mayalsohavearoleindevelopmentoftheblastocyst

(/embryology/index.php/File:Ova20he.jpg)
AntralFollicleandOocyte

Links:Zonapellucida(/embryology/index.php/Zona_pellucida)|MBoCFigure2021.Thezonapellucida
(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=mboc4.figgrp.3722)

CoronaRadiata
granulosacellsandextracellularmatrix
protectiveandnutritionalroleforcellsduringtransport
cellsarealsolostduringtransportalongoviduct

GameteformationSpermatogenesis
processofspermatagoniamatureintospermatazoa(sperm)
continuouslythroughoutlifeoccursintheseminiferoustubulesinthemalegonadtestis(pluraltestes)
atpubertyspermatagoniaactivateandproliferate(mitosis)
primaryspermatocyte>secondaryspermatocyte>spermatid>sperm
SeminiferousTubuleissiteofmaturationinvolvingmeiosisandspermiogenesis
SpermatogenesisMeiosis
meiosisisreductivecelldivision
1spermatagonia(diploid)46,XY(alsowritten44+XY)=4sperm(haploid)23,X23,X23,Y23,Y

(/embryology/index.php/File:Mouse
_zona_pellucida_01.jpg)
Mousezonapellucida[5]

Spermiogenesis
morphological(shape)changefromroundspermatidstoelongatedsperm
loosecytoplasm
Transformgolgiapparatusintoacrosome(inhead)
Organizemicrotubulesformotility(intail,flagellum)

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Segregatemitochondriaforenergy(intail)
Links:SpermatozoaDevelopment(/embryology/index.php/Spermatozoa_Development)

Ejaculate
HumanEjaculate
Byvolume<10%spermandaccessoryglandscontributemajorityofvolume(60%seminalvesicle,
10%bulbourethral,30%prostate)
3.5ml,200600millionsperm.
Capacitationistheremovalofglycoproteincoatandseminalproteinsandalterationofsperm
mitochondria.
Ovulationinducingfactoridentifiedinseveralspecies.
MarceloHRatto,YvonneALeduc,XimenaPValderrama,KarinEvanStraaten,LouisTJDelbaere,RogerA
Pierson,GreggPAdamsThenerveofovulationinducingfactorinsemen.Proc.Natl.Acad.Sci.U.S.A.:
2012,109(37)150427PubMed22908303(http://www.ncbi.nlm.nih.gov/pubmed/22908303)

(/embryology/index.php/File:Mouse_germinal_vesicle_03.jpg)
Mousegranulosacells[6]

InfertilitycanbeduetoOligospermia,Azoospermia,ImmotileCiliaSyndrome
Oligospermia(LowSpermCount)lessthan20millionspermafter72hourabstinencefromsex
Azoospermia(AbsentSperm)blockageofductnetwork
ImmotileCiliaSyndromelackofspermmotility
Links:Spermatozoa(/embryology/index.php/Spermatozoa_Development)|Prostate(/embryology/index.php/Prostate_Development)

FertilityWindow
Clinicalguidelineshavetypicallyidentifiedthe"fertilewindow"betweendays10and17withinthetypical28
daymenstrualcycle.
DatafromalargeUSANIEHSEarlyPregnancyStudy(/embryology/index.php/Reports#NIEHS_
_Early_Pregnancy_Study)(198286)identifiedthetimingofthefertilewindowwithinarangeofdifferent
menstrualcycles.[7]
fertilewindowoccurredduringabroadrangeofdaysinthemenstrualcycle.
betweendays6and21womenhadatminimuma10%probabilityofbeingintheirfertilewindow.
womencannotpredictasporadiclateovulation46%ofwomenwhosecycleshadnotyetresumedwere
potentiallyfertileinthefifthweekoftheircycle.
onlyabout30%ofwomenisthefertilewindowentirelywithinthedaysofthemenstrualcycleidentifiedby
clinicalguidelines(betweendays10and17)
mostwomenreachtheirfertilewindowearlierandothersmuchlater.
womenshouldbeadvisedthatthetimingoftheirfertilewindowcanbehighlyunpredictable,eveniftheir
cyclesareusuallyregular.

(/embryology/index.php/File:Menstrual_cycle_fertility_probabilit
Probabilityofwomenwithregularorirregular
cyclesbeingintheirfertilewindow

FertilizationSite
Fertilizationusuallyoccursinfirst1/3ofoviduct
Fertilizationcanalsooccuroutsideoviduct,associatedwithInVitroFertilization(IVF,GIFT,ZIFT...)and
ectopicpregnancy
Themajorityoffertilizedeggsdonotgoontoformanembryo

FertilizationSpermatozoa
SpermBindingzonapellucidaproteinZP2actsasreceptorforsperm
AcrosomeReactionexyocytosisofacrosomecontents(Calciummediated)MBoCFigure2031.The
acrosomereactionthatoccurswhenamammalianspermfertilizesanegg
(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mboc4.figgrp.3741)
enzymestodigestthezonapellucida
exposesspermsurfaceproteinstobindZP2
MembraneFusionbetweenspermandegg,allowsspermnucleipassageintoeggcytoplasm
Contactbetweenspermatozoaandoocyteeggcoat(zonapellucida[ZP])glycoproteinstriggersincreases
inintracellularcalciumion(iCa2+)concentrationinspermatozoa[8]
CATSPERchannelsonthedistalportionofsperm(theprincipalpiece)arerequiredfortheZPinduced
iCa2+increases
iCa2+increasestartsfromthespermatozoatailandpropagatestowardthehead
StoredepletionactivatedCa2+entryisthoughttomediatethesustainedphase

(/embryology/index.php/File:Week1_summary.jpg)
Week1

FertilizationOocyte
MembraneDepolarizationinnonmammalianspecies,causedbyspermmembranefusion,actsasaprimaryblocktopolyspermy.
CorticalReactionIP3pathwayelevatesintracellularCalcium,exocytosisofcorticalgranulesMBoCFigure2032.Howthecorticalreactioninamouse
eggisthoughttopreventadditionalspermfromenteringtheegg(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mboc4.figgrp.3743)
enzymealtersZP2soitwillnolongerbindspermplasmamembrane
Meiosis2completionof2ndmeioticdivision

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formssecondpolarbody(athirdpolarbodymaybeformedbymeioticdivisionofthefirstpolarbody)

SpermatozoaOocyteInteraction
MembraneFusion
Spermatozoaandoocytefusioninthemembraneadhesionarearequiresthepresenceof3membraneproteins(spermatozoaIzumo1oocytereceptorJunoand
Cd9).[9]

Izumo
ThespermspecificproteinIzumoisnamedforaJapaneseshrinededicatedtomarriageandisessentialforspermeggplasmamembranebindingandfusion.It
interactswiththespermatozoafolatereceptor4(Folr4).
Links:OMIM609278(http://omim.org/entry/609278)

Juno
folatereceptor4(Folr4)
Folatereceptorsarealsoknownasthefolicacid(FA)bindingproteinandbindof5methyltetrahydrofolate(5MeTHF).

CD9
Oocytecellsurfaceproteinactsasareceptorforpregnancyspecificglycoprotein17(Psg17).

Links:OMIM143030(http://omim.org/entry/143030)

Ovastacin

(AstacinLikeMetalloendopeptidase,ASTL)Anoocyteenzyme(zincdependentmetalloprotease)involvedin
alteringzonapellucidastructure,bycleavingzonapellucidaglycoprotein(ZP2),followingfertilisation.[10]This
changeinzonapellucidastructureisoneoftheblockerstopolyspermy.

(/embryology/index.php/File:Mouse_oocyte_cortical_granule_o
Mouseoocytecorticalgranule(green)
ovastacin(red)staining.[10]

FormationoftheZygote
PronucleiMaleandFemalehaploidnucleiapproacheachotherandnuclearmembranesbreakdown
chromosomalpairing,DNAreplicates,firstmitoticdivision
Spermatozoacontributescentriolewhichorganizesmitoticspindle
Oocytecontributesmitochondria(maternallyinherited)

Mitochondria
Mitochondriaofthespermatozoaarespecificallydestroyedinearlydevelopmentbyproteolysis(mouse4
to8celltransition).[11]
MetaphaseIIoocytesinratshaveanaveragemitochondrialDNA(mtDNA)copynumberof147,600
(+/3000)thatonlyincreasesatthe8cellstage.[12]

SexDetermination
baseduponwhetheranXorYcarryingspermhasfertilizedtheegg,shouldbe1.0sexratio.
actually1.05,105malesforevery100females,somestudiesshowmoremales2+daysafterovulation.
celltotipotent(equivalenttoastemcell,canformanytissueofthebody)
MenYChromosome

(/embryology/index.php/File:Early_zygote.jpg)
EarlyhumanzygoteshowingPronuclei

YChromosomecarriesSrygene,proteinproductactivatespathwayformalegonad(coveredingenitaldevelopment)

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WomenXChromosome
Genedosage,oneXchromosomeineachfemaleembryocellhastobeinactivated
processisapparentlyrandomandtherefore50%ofcellshavefather'sX,50%havemother'sX
Notethatbecausemenonlyhave1Xchromosome,ifabnormal,thisleadstoXlinkeddiseasesmorecommoninmalethatfemalewherebotheX'sneedto
beabnormal.

FertilizationProteinChanges
Arecentstudyinmicehasshownthatafterfertilizationthematernalproteinspresentintheoriginaloocytearequicklydegradedbythezygotestage.MIIoocytes
have185,643differentpeptideswhilezygotescontainonly85,369peptides.[13]

ProteinExpressionClassifiedbyMolecularFunctions
MIIoocyte

(/embryology/index.php/File:Mouse_MII_oocyte_protein_expression.jpg)

Zygote

(/embryology/index.php/File:Mouse_zygote_protein_expression.jpg)

Abnormalities
Theabnormalitieslistedbelowrelatetogeneticabnormalitiesoccurringduringfertilisation.Thereareofcourse
manyadditionalgeneticabnormalitiesinheritedandintroducedbytherecombinedmaternalandpaternal
genomes,otherthantrisomy21thesewillnotbecoveredhere.

CompleteHydatidiformMole
Chromosomalgeneticmaterialfromtheoocyte(ovum,egg)islost,byanunknownprocess.
Fertilizationthenoccurswithoneortwospermatozoaandanandrogenic(fromthemaleonly)conceptus
(fertilizedoocyte)isformed.
Theembryo(fetus,baby)doesnotdevelopatallbuttheplacentadoesgrow.
Links:AbnormalDevelopmentHydatidiformMole(/embryology/index.php/Abnormal_Development_
_Hydatidiform_Mole)

PartialHydatidiformMole
Threesetsofchromosomesinsteadoftheusualtwoandthisiscalledtriploidy.
chromosomal(genetic)materialfromtheoocyte(ovum,egg)isretainedandtheeggisfertilizedbyoneor
twospermatozoa.
withpartialmoletherearematernalchromosomesandthereisafetus.
thethreesetsofchromosomesmeansthefetusisalwaysgrosslyabnormalandwillnotsurvive.

(/embryology/index.php/Trisomy_21)
Trisomy21

Links:AbnormalDevelopmentHydatidiformMole(/embryology/index.php/Abnormal_Development__Hydatidiform_Mole)

Parthenogenesis
(Greek,parthenos=virgin,genesis=birth)Thedevelopmentofanunfertilizedoocyte(nospermatozoa).Otherthanmammals,manydifferentspecies(plants,
insects,reptiles)candevelopfromunfertilizedeggs.Anembryosoformedwithoutspermcontribution.Blockingofparthenogenesisinmammalsappearstobe
relatedtogenomicimprinting.Abnormalparthenogenicprocessescanoccurinmammals,andmorerecentlyaparthenogenicmousehasbeenmadeinthe
laboratory.

Trisomy21
Trisomy21(Down'sorDownsyndrome)iscausedbynondisjunctionofchromosome21inaparentwhoischromosomallynormalandisoneofthemost
commonchromosomalaneuploidyabnormalitiesinlivebornchildren.Thefrequencyoftrisomy21inthepopulationisapproximately1in650to1,000livebirths,
inAustraliabetween199197therewere2,358Trisomy21infants.Thereareotherlessfrequentlyoccurringtrisomies(18,13andX).

Links:Trisomy21(/embryology/index.php/Trisomy_21)

References
1.DiegoRafaelUates,HctorAlejandroGuidobaldi,LauraVirginiaGatica,MarisaAnglicaCubilla,MaraEugeniaTeves,AyelnMoreno,LauraCecilia
GiojalasVersatileactionofpicomolargradientsofprogesteroneondifferentspermsubpopulations.PLoSONE:2014,9(3)e91181PubMed

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24614230(http://www.ncbi.nlm.nih.gov/pubmed/24614230)
2.YasuhiroYamauchi,JeffreyAShaman,WStevenWardNongeneticcontributionsofthespermnucleustoembryonicdevelopment.AsianJ.Androl.:
2011,13(1)315PubMed20953203(http://www.ncbi.nlm.nih.gov/pubmed/20953203)
3.AntoineJgou,AhmedZiyyat,VirginieBarraudLange,EricPerez,JeanPhilippeWolf,FrdricPincet,ChristineGourierCD9tetraspaningeneratesfusion
competentsitesontheeggmembraneformammalianfertilization.Proc.Natl.Acad.Sci.U.S.A.:2011,108(27)1094651PubMed21690351
(http://www.ncbi.nlm.nih.gov/pubmed/21690351)
4.GagandeepGahlay,LynGauthier,BorisBaibakov,OlgaEpifano,JurrienDeanGameterecognitioninmicedependsonthecleavagestatusofanegg's
zonapellucidaprotein.Science:2010,329(5988)2169PubMed20616279(http://www.ncbi.nlm.nih.gov/pubmed/20616279)
5.PaulMWassarmanZonapellucidaglycoproteins.J.Biol.Chem.:2008,283(36)242859PubMed18539589
(http://www.ncbi.nlm.nih.gov/pubmed/18539589)|JBiolChem.(http://www.jbc.org/content/283/36/24285.long)
6.HongXiaZhou,YuZhenMa,YingLeiLiu,YingChen,ChengJieZhou,ShaNaWu,JiangPengShen,ChengGuangLiangAssessmentofmouse
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Textbooks
HumanEmbryology(2nded.)LarsonCh1p132
TheDevelopingHuman:ClinicallyOrientedEmbryology(6thed.)MooreandPersaud
BeforeweAreBorn(5thed.)MooreandPersaudCh2p1433
EssentialsofHumanEmbryologyLarsonCh1p116
HumanEmbryologyFitzgeraldandFitzgeraldCh2p814
SearchNCBIBookshelffertilization(http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=fertilization)|fertilisation
(http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=fertilisation)

Reviews
IJDBVol.52Nos.5/6(2008)Fertilization(http://www.ijdb.ehu.es/web/contents.php?vol=52&issue=56)

Articles
AJWilcox,CRWeinberg,DDBairdPostovulatoryageingofthehumanoocyteandembryofailure.Hum.Reprod.:1998,13(2)3947PubMed9557845
(http://www.ncbi.nlm.nih.gov/pubmed/9557845)
|HumReprod.(http://humrep.oxfordjournals.org/content/13/2/394.abstract)

SearchPubmed
April2010
fertilizationAll(51803)Review(5928)FreeFullText(11715)
SearchPubmedNow:fertilization(http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=fertilization)|fertilisation
(http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=fertilisation)|zonapellucida(http://www.ncbi.nlm.nih.gov/sites/entrez?
db=pubmed&cmd=search&term=zona+pellucida)|zygote(http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=zygote)

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