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(/embryology/index.php/File:Human_fertilization_movie_2_frame_03.jpg)Human Fertilization

Fertilization

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Contents1 Introduction2 Some Recent Findings3 Objectives4 Movies5 Fertilization Preparation6 Oogenesis7 Menstrual Cycle8 Ovulation (HPG Axis)9 Zona Pellucida10 Corona Radiata11 Gamete formation­ Spermatogenesis

11.1 Spermiogenesis12 Ejaculate13 Fertility Window14 Fertilization Site

14.1 Fertilization ­ Spermatozoa14.2 Fertilization ­ Oocyte

15 Spermatozoa ­ Oocyte Interaction15.1 Membrane Fusion15.2 Izumo15.3 Juno15.4 CD915.5 Ovastacin

16 Formation of the Zygote16.1 Mitochondria

17 Sex Determination18 Fertilization Protein Changes

18.1 Protein Expression Classified by Molecular Functions19 Abnormalities

19.1 Complete Hydatidiform Mole19.2 Partial Hydatidiform Mole19.3 Parthenogenesis19.4 Trisomy 21

20 References20.1 Textbooks20.2 Reviews20.3 Articles20.4 Search Pubmed

21 Additional Images22 External Links23 Glossary Links

IntroductionFertilization is the fusion of haploid gametes, egg and sperm, to form the diploid zygote. Note though there canbe subtle differences in the fertilization process which occurs naturally within the body or through reproductivetechnologies outside the body, the overall product in both cases is a diplod zygote. In fertilization research,after humans the mouse is the most studied species followed by domestic and farm animals. The process offertilization involves components of, and signaling between, both sperm (spermatozoa) and egg (oocyte).

In addition to in vivo fertilization there are many new in vitro technologies related to human infertility (AssistedReproductive Technology (/embryology/index.php/Assisted_Reproductive_Technology)) and animal productionsomatic cell nuclear transfer (SCNT) to generate a zygote.

Note different spelling ­ USA spelling "Fertilization", Australian spelling "Fertilisation".

Fertilization Links: Fertilization | Oocyte (/embryology/index.php/Oocyte_Development) | Spermatozoa(/embryology/index.php/Spermatozoa_Development) | Cell Division ­ Meiosis(/embryology/index.php/Cell_Division_­_Meiosis) | Zona pellucida (/embryology/index.php/Zona_pellucida)| Zygote (/embryology/index.php/Zygote) | Lecture ­ Fertilization (/embryology/index.php/Lecture_­_Fertilization) | Cell Division ­ Mitosis(/embryology/index.php/Cell_Division_­_Mitosis) | Lecture ­ Week 1 and 2 (/embryology/index.php/Lecture_­_Week_1_and_2_Development) | Hydatidiform

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(/embryology/index.php/File:Early_zygote_labelled.jpg)Early Human Zygote

(/embryology/index.php/File:Mouse­fertilization_01.jpg)The first polar body deforms the mammalianegg away from its encapsulating zonapellucida

Mole (/embryology/index.php/Abnormal_Development_­_Hydatidiform_Mole) | Assisted ReproductiveTechnology (/embryology/index.php/Assisted_Reproductive_Technology) | Lecture ­ Genital Development(/embryology/index.php/Lecture_­_Genital_Development)| Menstrual Cycle(/embryology/index.php/Menstrual_Cycle) | Morula (/embryology/index.php/Morula) | Blastocyst(/embryology/index.php/Blastocyst)

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

Some Recent FindingsVersatile action of picomolar gradients of progesterone on different sperm subpopulations "Highstep concentrations of progesterone may stimulate various sperm physiological processes, such aspriming and the acrosome reaction. However, approaching the egg, spermatozoa face increasingconcentrations of the hormone, as it is secreted by the cumulus cells and then passively diffuses along thecumulus matrix and beyond. ... The results suggest a versatile role of the gradual distribution of very lowdoses of progesterone, which selectively stimulate the priming and the acrosome reaction in differentsperm subpopulations."Juno is the egg Izumo receptor and is essential for mammalian fertilization "Fertilization occurswhen sperm and egg recognize each other and fuse to form a new, genetically distinct organism. Themolecular basis of sperm–egg recognition is unknown, but is likely to require interactions between receptorproteins displayed on their surface. Izumo1 is an essential sperm cell­surface protein, but its receptor onthe egg has not been described. Here we identify folate receptor 4 (Folr4) as the receptor for Izumo1 onthe mouse egg, and propose to rename it Juno. We show that the Izumo1–Juno interaction is conservedwithin several mammalian species, including humans. Female mice lacking Juno are infertile and Juno­deficient eggs do not fuse with normal sperm. Rapid shedding of Juno from the oolemma after fertilizationsuggests a mechanism for the membrane block to polyspermy, ensuring eggs normally fuse with just asingle sperm."Nature 16 April 2014(http://www.nature.com/nature/journal/vaop/ncurrent/full/nature13203.html)Non­genetic contributions of the sperm nucleus to embryonic development "Recent data fromseveral laboratories have provided evidence that the newly fertilized oocyte inherits epigenetic signalsfrom the sperm chromatin that are required for proper embryonic development. For the purposes of thisreview, the term epigenetic is used to describe all types of molecular information that are transmitted fromthe sperm cell to the embryo. There are at least six different forms of epigenetic information that havealready been established as being required for proper embryogenesis in mammals or for which there isevidence that it may do so. These are (i) DNA methylation; (ii) sperm­specific histones, (iii) otherchromatin­associated proteins; (iv) the perinuclear theca proteins; (v) sperm­born RNAs and, the focus ofthis review; and (vi) the DNA loop domain organization by the sperm nuclear matrix. These epigeneticsignals should be considered when designing protocols for the manipulation and cryopreservation ofspermatozoa for assisted reproductive technology as necessary components for effective fertilization andsubsequent embryo development."CD9 tetraspanin generates fusion competent sites on the egg membrane for mammalianfertilization "CD9 tetraspanin is the only egg membrane protein known to be essential for fertilization.To investigate its role, we have measured, on a unique acrosome reacted sperm brought in contact withan egg, the adhesion probability and strength with a sensitivity of a single molecule attachment. Probingthe binding events at different locations of wild­type egg we described different modes of interaction. Here,we show that more gamete adhesion events occur on Cd9 null eggs but that the strongest interactionmode disappears. We propose that sperm­egg fusion is a direct consequence of CD9 controlled sperm­egg adhesion properties. CD9 generates adhesion sites responsible for the strongest of the observedgamete interaction. These strong adhesion sites impose, during the whole interaction lifetime, a tightproximity of the gamete membranes, which is a requirement for fusion to take place. The CD9­inducedadhesion sites would be the actual location where fusion occurs."Gamete recognition in mice depends on the cleavage status of an egg's zona pellucidaprotein "sperm­egg recognition depends on the cleavage status of ZP2 and that binding at the surface ofthe zona is not sufficient to induce sperm acrosome exocytosis."

More recent papers

ObjectivesUnderstand the mechanisms of gamete formation.Understand the mechanisms of cell division.Describe the differences between mitosis and meiosis.Understand the mechanisms of fertilization, both in vivo and in vitro.Describe the cleavage of the zygote.Have a preliminary understanding of the role and process in male sex determination and X inactivation.Understand the abnormalities that occur during this period of development.

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(/embryology/index.php/File:Ovary5x.gif)Histology of the Ovary

Movies

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

(/embryology/index.php/Spermatozoa_Structure_Movie) SpermatozoaPage(/embryology/index.php/Spermatozoa_Structure_Movie)| Play(/embryology/images/5/5b/Spermatozoa_animation.mp4)

(/embryology/index.php/Human_Spermatozoa_Motility_Movie) Spermatozoa MotilityPage(/embryology/index.php/Human_Spermatozoa_Motility_Movie)| Play(/embryology/images/a/ac/Spermatozoa_motility_01.mp4)

(/embryology/index.php/Human_Fertilization_Movie) Page(/embryology/index.php/Human_Fertilization_Movie)Play(/embryology/images/6/6c/Human_fertilization_01.mp4)

(/embryology/index.php/Fertilization_Movie) FertilizationPage(/embryology/index.php/Fertilization_Movie) |Play(/embryology/images/5/5f/Fertilization_003.mp4)

(/embryology/index.php/Mouse_Fertilization_Movie) Mouse FertilisationPage(/embryology/index.php/Mouse_Fertilization_Movie) |Play (/embryology/images/c/c0/Fertilization_001.mp4)

(/embryology/index.php/Pronuclear_Fusion_Movie) Pronuclear FusionPage (/embryology/index.php/Pronuclear_Fusion_Movie) |Play (/embryology/images/9/95/Pronuclear_fusion_001.mp4)

(/embryology/index.php/Week_1_Movie) Page(/embryology/images/b/b4/Week1_001.mp4)

Fertilization PreparationPrior to the fertilization process commencing both the gametes oocyte (egg) and spermatozoa (sperm) require completion of a number of biological processes.

Oocyte Meiosis ­ completes Meiosis 1 and commences Meiosis 2 (arrests at Metaphase II).Spermatozoa Capacitation ­ following release (ejaculation) and mixing with other glandular secretions, activates motility and acrosome preparation.Migration ­ both Oocyte and Spermatozoa.

oocyte ovulation and release with associated cells, from ovary into fimbria then into uterine tube (oviduct, uterine horn, fallopian tube) and epithelial ciliamediated movement.spermatozoa ejaculation, deposited in vagina, movement of tail to "swim" in uterine secretions through cervix, uterine body and into uterine tube, haveapproximately 24­72h to fertilize oocyte.

Endocrinology ­ Diagram of the comparative anatomy of the male and female reproductive tracts (http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=endocrin&part=A972&rendertype=box&id=A1230)

OogenesisProcess of oogonia mature into oocytes (ova, ovum, egg)all oogonia form primary oocytes before birth, therefore a maturation of preexisting cells in the femalegonad, ovary

(/embryology/index.php/File:Human_ovary_non­

growing_follicle_model.jpg)

humans usually only 1 ovum released every menstrual cycle (IVF­ superovulation)oocyte and its surrounding cells = follicleprimary ­> secondary ­> ovulation releases

Ovary­ Histology ­ whole transverse section (cortex, medulla)

Menstrual CyclePrimary Oocyte ­ arrested at early Meiosis 1

diploid: 22 chromosome pairs + 1 pair X chromosomes (46, XX)autosomes and sex chromosome

Oogenesis­ pre­antral then antral follicle (Graafian follicle is mature antral follicle released)Secondary oocyte

1 Day before ovulation completes (stim by LH) Meiosis 1haploid: 22 chromosomes + 1 X chromosome (23, X)nondisjunction­ abnormal chromosome segregationbegins Meiosis 2 and arrests at metaphasenote no interphase replication of DNA, only fertilization will complete Meiosis 2

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(/embryology/index.php/File:Ova41he.jpg)Preantral Follicle

(/embryology/index.php/File:Ova20he.jpg)Antral Follicle and Oocyte

(/embryology/index.php/File:Mouse­_zona_pellucida_01.jpg)Mouse zona pellucida

Ovulation (HPG Axis)

(/embryology/index.php/File:Menstrual_cycle.png)

Hypothalmus releases gonadotropin releasing hormone (GRH, luteinizing hormone–releasing hormone,LHRH) ­> Pituitary releases follicle stimulating hormone (FSH) and lutenizing hormone (LH) ­> ovary follicledevelopment and ovulation.

release of the secondary oocyte and formation of corpus luteumsecondary oocyte encased in zona pellucida and corona radiata

Ovulation associated with follicle rupture and ampulla movement.

Zona Pellucidaglycoprotein shell ZP1, ZP2, ZP3, ZP4 (mouse only ZP1­3)mechanical protection of egginvolved in the fertilization processsperm bindingadhesion of sperm to eggacrosome reaction

releases enzymes to locally breakdownblock of polyspermy

altered to prevent more than 1 sperm penetratingmay also have a role in development of the blastocyst

Links: Zona pellucida (/embryology/index.php/Zona_pellucida) | MBoC ­ Figure 20­21. The zona pellucida(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=mboc4.figgrp.3722)

Corona Radiatagranulosa cells and extracellular matrixprotective and nutritional role for cells during transportcells are also lost during transport along oviduct

Gamete formation­ Spermatogenesisprocess of spermatagonia mature into spermatazoa (sperm)continuously throughout life occurs in the seminiferous tubules in the male gonad­ testis (plural testes)at puberty spermatagonia activate and proliferate (mitosis)primary spermatocyte ­> secondary spermatocyte­> spermatid­>spermSeminiferous Tubule is site of maturation involving meiosis and spermiogenesisSpermatogenesis­ Meiosismeiosis is reductive cell division

1 spermatagonia (diploid) 46, XY (also written 44+XY) = 4 sperm (haploid); 23, X 23, X 23, Y 23, Y

Spermiogenesismorphological (shape) change from round spermatids to elongated spermloose cytoplasmTransform golgi apparatus into acrosome (in head)Organize microtubules for motility (in tail, flagellum)

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(/embryology/index.php/File:Mouse_germinal_vesicle_03.jpg)Mouse granulosa cells

(/embryology/index.php/File:Menstrual_cycle_fertility_probability_01.jpg)Probability of women with regular or irregularcycles being in their fertile window

(/embryology/index.php/File:Week1_summary.jpg)Week 1

Segregate mitochondria for energy (in tail)

Links: Spermatozoa Development (/embryology/index.php/Spermatozoa_Development)

EjaculateHuman Ejaculate

By volume <10 % sperm and accessory glands contribute majority of volume (60 % seminal vesicle,10 % bulbourethral, 30 % prostate)3.5 ml, 200­600 million sperm.

Capacitation is the removal of glycoprotein coat and seminal proteins and alteration of spermmitochondria.Ovulation­inducing factor identified in several species.

Marcelo H Ratto, Yvonne A Leduc, Ximena P Valderrama, Karin E van Straaten, Louis T J Delbaere, Roger APierson, Gregg P Adams The nerve of ovulation­inducing factor in semen. Proc. Natl. Acad. Sci. U.S.A.:2012, 109(37);15042­7 PubMed 22908303 (http://www.ncbi.nlm.nih.gov/pubmed/22908303)

Infertility can be due to Oligospermia, Azoospermia, Immotile Cilia SyndromeOligospermia (Low Sperm Count) ­ less than 20 million sperm after 72 hour abstinence from sexAzoospermia (Absent Sperm) ­ blockage of duct networkImmotile Cilia Syndrome ­ lack of sperm motility

Links: Spermatozoa (/embryology/index.php/Spermatozoa_Development) | Prostate (/embryology/index.php/Prostate_Development)

Fertility WindowClinical guidelines have typically identified the "fertile window" between days 10 and 17 within the typical 28day menstrual cycle.

Data from a large USA NIEHS ­ Early Pregnancy Study (/embryology/index.php/Reports#NIEHS_­_Early_Pregnancy_Study) (1982­86) identified the timing of the “fertile window” within a range of differentmenstrual cycles.

fertile window occurred during a broad range of days in the menstrual cycle.between days 6 and 21 women had at minimum a 10% probability of being in their fertile window.women cannot predict a sporadic late ovulation; 4 ­ 6% of women whose cycles had not yet resumed werepotentially fertile in the fifth week of their cycle.only about 30% of women is the fertile window entirely within the days of the menstrual cycle identified byclinical guidelines (between days 10 and 17)most women reach their fertile window earlier and others much later.women should be advised that the timing of their fertile window can be highly unpredictable, even if theircycles are usually regular.

Fertilization SiteFertilization usually occurs in first 1/3 of oviductFertilization can also occur outside oviduct, associated with In Vitro Fertilization (IVF, GIFT, ZIFT...) andectopic pregnancyThe majority of fertilized eggs do not go on to form an embryo

Fertilization ­ SpermatozoaSperm Binding ­ zona pellucida protein ZP2 acts as receptor for spermAcrosome Reaction ­ exyocytosis of acrosome contents (Calcium mediated) MBoC ­ Figure 20­31. Theacrosome reaction that occurs when a mammalian sperm fertilizes an egg(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mboc4.figgrp.3741)

enzymes to digest the zona pellucidaexposes sperm surface proteins to bind ZP2

Membrane Fusion ­ between sperm and egg, allows sperm nuclei passage into egg cytoplasmContact between spermatozoa and oocyte egg coat (zona pellucida [ZP]) glycoproteins triggers increasesin intracellular calcium ion (iCa ) concentration in spermatozoaCATSPER channels on the distal portion of sperm (the principal piece) are required for the ZP­inducediCa increasesiCa increase starts from the spermatozoa tail and propagates toward the headStore depletion­activated Ca entry is thought to mediate the sustained phase

Fertilization ­ OocyteMembrane Depolarization ­ in non­mammalian species, caused by sperm membrane fusion, acts as a primary block to polyspermy.Cortical Reaction ­ IP3 pathway elevates intracellular Calcium, exocytosis of cortical granules MBoC ­ Figure 20­32. How the cortical reaction in a mouseegg is thought to prevent additional sperm from entering the egg (http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mboc4.figgrp.3743)

enzyme alters ZP2 so it will no longer bind sperm plasma membraneMeiosis 2 ­ completion of 2nd meiotic division

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(/embryology/index.php/File:Mouse_oocyte_cortical_granule_ovastacin_01.jpg)Mouse oocyte cortical granule (green)ovastacin (red) staining.

(/embryology/index.php/File:Early_zygote.jpg)Early human zygote showing Pronuclei

forms second polar body (a third polar body may be formed by meiotic division of the first polar body)

Spermatozoa ­ Oocyte InteractionMembrane FusionSpermatozoa and oocyte fusion in the membrane adhesion area requires the presence of 3 membrane proteins (spermatozoa Izumo1; oocyte receptor Juno andCd9).

IzumoThe sperm­specific protein Izumo is named for a Japanese shrine dedicated to marriage and is essential for sperm­egg plasma membrane binding and fusion. Itinteracts with the spermatozoa folate receptor 4 (Folr4).

Links: OMIM 609278 (http://omim.org/entry/609278)

Junofolate receptor 4 (Folr4)

Folate receptors are also known as the folic acid (FA) binding protein and bind of 5­methyltetrahydrofolate (5­MeTHF).

CD9Oocyte cell surface protein acts as a receptor for pregnancy­specific glycoprotein 17 (Psg17).

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

Ovastacin

(Astacin­Like Metalloendopeptidase, ASTL) An oocyte enzyme (zinc­dependent metalloprotease) involved inaltering zona pellucida structure, by cleaving zona pellucida glycoprotein (ZP2), following fertilisation. Thischange in zona pellucida structure is one of the blockers to polyspermy.

Formation of the ZygotePronuclei ­ Male and Female haploid nuclei approach each other and nuclear membranes break downchromosomal pairing, DNA replicates, first mitotic divisionSpermatozoa contributes ­ centriole which organizes mitotic spindleOocyte contributes ­ mitochondria (maternally inherited)

MitochondriaMitochondria of the spermatozoa are specifically destroyed in early development by proteolysis (mouse 4to 8 cell transition).Metaphase II oocytes in rats have an average mitochondrial DNA (mtDNA) copy number of 147,600(+/­3000) that only increases at the 8­cell stage.

Sex Determinationbased upon whether an X or Y carrying sperm has fertilized the egg, should be 1.0 sex ratio.actually 1.05, 105 males for every 100 females, some studies show more males 2+ days after ovulation.cell totipotent (equivalent to a stem cell, can form any tissue of the body)

Men ­ Y Chromosome

Y Chromosome carries Sry gene, protein product activates pathway for male gonad (covered in genital development)

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(/embryology/index.php/Trisomy_21)Trisomy 21

Women ­ X Chromosome

Gene dosage, one X chromosome in each female embryo cell has to be inactivatedprocess is apparently random and therefore 50% of cells have father's X, 50% have mother's XNote that because men only have 1 X chromosome, if abnormal, this leads to X­linked diseases more common in male that female where bothe X's need tobe abnormal.

Fertilization Protein ChangesA recent study in mice has shown that after fertilization the maternal proteins present in the original oocyte are quickly degraded by the zygote stage. MII oocyteshave 185,643 different peptides while zygotes contain only 85,369 peptides.

Protein Expression Classified by Molecular FunctionsMII oocyte Zygote

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

AbnormalitiesThe abnormalities listed below relate to genetic abnormalities occurring during fertilisation. There are of coursemany additional genetic abnormalities inherited and introduced by the recombined maternal and paternalgenomes, other than trisomy 21 these will not be covered here.

Complete Hydatidiform MoleChromosomal genetic material from the oocyte (ovum, egg) is lost, by an unknown process.Fertilization then occurs with one or two spermatozoa and an androgenic (from the male only) conceptus(fertilized oocyte) is formed.The embryo (fetus, baby) does not develop at all but the placenta does grow.Links: Abnormal Development ­ Hydatidiform Mole (/embryology/index.php/Abnormal_Development_­_Hydatidiform_Mole)

Partial Hydatidiform MoleThree sets of chromosomes instead of the usual two and this is called triploidy.

chromosomal (genetic) material from the oocyte (ovum, egg) is retained and the egg is fertilized by one ortwo spermatozoa.with partial mole there are maternal chromosomes and there is a fetus.the three sets of chromosomes means the fetus is always grossly abnormal and will not survive.Links: Abnormal Development ­ Hydatidiform Mole (/embryology/index.php/Abnormal_Development_­_Hydatidiform_Mole)

Parthenogenesis(Greek, parthenos = virgin, genesis = birth) The development of an unfertilized oocyte (no spermatozoa). Other than mammals, many different species (plants,insects, reptiles) can develop from unfertilized eggs. An embryo so formed without sperm contribution. Blocking of parthenogenesis in mammals appears to berelated to genomic imprinting. Abnormal parthenogenic processes can occur in mammals, and more recently a parthenogenic mouse has been made in thelaboratory.

Trisomy 21Trisomy 21 (Down's or Down syndrome) is caused by nondisjunction of chromosome 21 in a parent who is chromosomally normal and is one of the mostcommon chromosomal aneuploidy abnormalities in liveborn children. The frequency of trisomy 21 in the population is approximately 1 in 650 to 1,000 live births,in Australia between 1991­97 there were 2,358 Trisomy 21 infants. There are other less frequently occurring trisomies (18, 13 and X).

Links: Trisomy 21 (/embryology/index.php/Trisomy_21)

References1. Diego Rafael Uñates, Héctor Alejandro Guidobaldi, Laura Virginia Gatica, Marisa Angélica Cubilla, María Eugenia Teves, Ayelén Moreno, Laura Cecilia

Giojalas Versatile action of picomolar gradients of progesterone on different sperm subpopulations. PLoS ONE: 2014, 9(3);e91181 PubMed

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24614230 (http://www.ncbi.nlm.nih.gov/pubmed/24614230)2. Yasuhiro Yamauchi, Jeffrey A Shaman, W Steven Ward Non­genetic contributions of the sperm nucleus to embryonic development. Asian J. Androl.:

2011, 13(1);31­5 PubMed 20953203 (http://www.ncbi.nlm.nih.gov/pubmed/20953203)3. Antoine Jégou, Ahmed Ziyyat, Virginie Barraud­Lange, Eric Perez, Jean Philippe Wolf, Frédéric Pincet, Christine Gourier CD9 tetraspanin generates fusion

competent sites on the egg membrane for mammalian fertilization. Proc. Natl. Acad. Sci. U.S.A.: 2011, 108(27);10946­51 PubMed 21690351(http://www.ncbi.nlm.nih.gov/pubmed/21690351)

4. Gagandeep Gahlay, Lyn Gauthier, Boris Baibakov, Olga Epifano, Jurrien Dean Gamete recognition in mice depends on the cleavage status of an egg'szona pellucida protein. Science: 2010, 329(5988);216­9 PubMed 20616279 (http://www.ncbi.nlm.nih.gov/pubmed/20616279)

5. Paul M Wassarman Zona pellucida glycoproteins. J. Biol. Chem.: 2008, 283(36);24285­9 PubMed 18539589(http://www.ncbi.nlm.nih.gov/pubmed/18539589) | J Biol Chem. (http://www.jbc.org/content/283/36/24285.long)

6. Hong­Xia Zhou, Yu­Zhen Ma, Ying­Lei Liu, Ying Chen, Cheng­Jie Zhou, Sha­Na Wu, Jiang­Peng Shen, Cheng­Guang Liang Assessment of mousegerminal vesicle stage oocyte quality by evaluating the cumulus layer, zona pellucida, and perivitelline space. PLoS ONE: 2014, 9(8);e105812PubMed 25144310 (http://www.ncbi.nlm.nih.gov/pubmed/25144310) | PLoS One.(http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0105812)

7. A J Wilcox, D Dunson, D D Baird The timing of the "fertile window" in the menstrual cycle: day specific estimates from a prospective study. BMJ:2000, 321(7271);1259­62 PubMed 11082086 (http://www.ncbi.nlm.nih.gov/pubmed/11082086) | PMC27529(http://www.ncbi.nlm.nih.gov/pmc/articles/PMC27529) | BMJ (http://www.bmj.com/content/321/7271/1259?view=long&pmid=11082086)

8. Jingsheng Xia, Dejian Ren Egg coat proteins activate calcium entry into mouse sperm via CATSPER channels. Biol. Reprod.: 2009, 80(6);1092­8PubMed 19211808 (http://www.ncbi.nlm.nih.gov/pubmed/19211808)

9. Myriam Chalbi, Virginie Barraud­Lange, Benjamin Ravaux, Kevin Howan, Nicolas Rodriguez, Pierre Soule, Arnaud Ndzoudi, Claude Boucheix, EricRubinstein, Jean Philippe Wolf, Ahmed Ziyyat, Eric Perez, Frédéric Pincet, Christine Gourier Binding of sperm protein Izumo1 and its egg receptor Junodrives Cd9 accumulation in the intercellular contact area prior to fusion during mammalian fertilization. Development: 2014, 141(19);3732­9 PubMed25209248 (http://www.ncbi.nlm.nih.gov/pubmed/25209248)

10. Anna D Burkart, Bo Xiong, Boris Baibakov, Maria Jiménez­Movilla, Jurrien Dean Ovastacin, a cortical granule protease, cleaves ZP2 in the zonapellucida to prevent polyspermy. J. Cell Biol.: 2012, 197(1);37­44 PubMed 22472438 (http://www.ncbi.nlm.nih.gov/pubmed/22472438)

11. J M Cummins Fertilization and elimination of the paternal mitochondrial genome. Hum. Reprod.: 2000, 15 Suppl 2;92­101 PubMed 11041517(http://www.ncbi.nlm.nih.gov/pubmed/11041517)

12. Yuichi Kameyama, France Filion, Jae Gyu Yoo, Lawrence C Smith Characterization of mitochondrial replication and transcription control during ratearly development in vivo and in vitro. Reproduction: 2007, 133(2);423­32 PubMed 17307910 (http://www.ncbi.nlm.nih.gov/pubmed/17307910)

13. Shufang Wang, Zhaohui Kou, Zhiyi Jing, Yu Zhang, Xinzheng Guo, Mengqiu Dong, Ian Wilmut, Shaorong Gao Proteome of mouse oocytes at differentdevelopmental stages. Proc. Natl. Acad. Sci. U.S.A.: 2010, 107(41);17639­44 PubMed 20876089 (http://www.ncbi.nlm.nih.gov/pubmed/20876089) | PNAS(http://www.pnas.org/content/107/41/17639.full)

TextbooksHuman Embryology (2nd ed.) Larson Ch1 p1­32The Developing Human: Clinically Oriented Embryology (6th ed.) Moore and PersaudBefore we Are Born (5th ed.) Moore and Persaud Ch 2 p14­33Essentials of Human Embryology Larson Ch1 p1­16Human Embryology Fitzgerald and Fitzgerald Ch2 p8­14

Search NCBI Bookshelf fertilization (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)

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

ArticlesA J Wilcox, C R Weinberg, D D Baird Post­ovulatory ageing of the human oocyte and embryo failure. Hum. Reprod.: 1998, 13(2);394­7 PubMed 9557845(http://www.ncbi.nlm.nih.gov/pubmed/9557845)

| Hum Reprod. (http://humrep.oxfordjournals.org/content/13/2/394.abstract)

Search PubmedApril 2010

fertilization ­ All (51803) Review (5928) Free Full Text (11715)

Search Pubmed Now: 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) | zona pellucida (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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4/6/2016 Fertilization ­ Embryology

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Mouse ­ Oocyte polarbody

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Mouse ­ SpermatozoaSwims within thePrevitelline space (PVS)Prior to Fertilization

External LinksExternal Links Notice ­ The dynamic nature of the internet may mean that some of these listed links may no longer function. If the link no longer works searchthe web with the link text or name.

PBS ­ First Human Eggs Fertilized in a Laboratory (http://www.pbs.org/wgbh/americanexperience/features/general­article/babies­first­eggs­fertilized) 1944.

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Cite this page: Hill, M.A. (2016) Embryology Fertilization. Retrieved April 5, 2016, from https://embryology.med.unsw.edu.au/embryology/index.php/Fertilization(https://embryology.med.unsw.edu.au/embryology/index.php/Fertilization)

What Links Here? (http://php.med.unsw.edu.au/embryology/index.php?title=Special:WhatLinksHere/Fertilization)

© Dr Mark Hill 2016, UNSW Embryology ISBN: 978 0 7334 2609 4 ­ UNSW CRICOS Provider Code No. 00098G

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