Z8600021: Blanked the page

2019-04-15T11:02:45Z

Blanked the page

Show changes

Z8600021: Undo revision 376941 by Z8600021 (talk)

2019-04-15T11:01:59Z

Undo revision 376941 by Z8600021 (talk)

Show changes

Z8600021: Blanked the page

2019-04-15T11:01:38Z

Blanked the page

Show changes

Z8600021 at 10:59, 15 April 2019

2019-04-15T10:59:00Z

← Older revision		Revision as of 21:59, 15 April 2019
Line 1:		Line 1:

	{{Header}}		{{Header}}
	==Introduction==		==Introduction==

Z8600021: Created page with " {{Header}} ==Introduction== Oocyte and developing zona pellucida in the ovary I..."

2019-04-15T10:58:14Z

Created page with " {{Header}} ==Introduction== thumb|alt=Oocyte and developing zona pellucida in the ovary|Oocyte and developing zona pellucida in the ovary I..."

New page

{{Header}}
==Introduction==
[[File:Ovary histology 061.jpg|thumb|alt=Oocyte and developing zona pellucida in the ovary|Oocyte and developing zona pellucida in the ovary]]
[[Image:CSt3.jpg|thumb|alt=Human blastocyst "hatching" from the zona pellucida|Human blastocyst "hatching" from the zona pellucida ([[Week 1|week 1]] [[Carnegie_stage_3|Carnegie stage 3]])]]

(Latin, ''zona pellucida'' = transparent zone) The {{zona pellucida}} (ZP) is a specialized extracellular matrix surrounding the developing oocyte (egg, ovum) within each follicle within the ovary. This thick matrix is thought to be formed by secretions from the oocyte and the follicle granulosa cells and in human oocytes consists of four types of zona pellucida glycoproteins {{ZP1}}, {{ZP2}}, {{ZP3}} and {{ZP4}} which have different roles in fertilization. Note that mice also have 4 ZP genes, but multiple stop and missense codons in ZP4 means that only three are produced. Polymers of ZP2 and ZP3 organized into extended filaments that are cross-linked by ZP1 homodimers.

Note that depending upon species and type of study, the zona pellucida can also be called the: oolemma, egg coat or vitelline membrane.

The zona pellucid has many different roles including in oocyte development, protection during growth and transport, {{fertilization}}, {{spermatozoa}} binding, preventing [[P#polyspermy|polyspermy]], {{blastocyst}} development, and preventing premature implantation ({{ectopic pregnancy}}).

{|
| [[File:Human_oocyte_11.jpg|300px]]
| [[File:Early_zygote.jpg|350px]]
|-
| Human oocyte contained inside zona pellucida.{{#pmid:24586757|PMID24586757}}
| Early zygote inside zona pellucida
|}
In human development, during the first week of development following fertilization the zona pellucida remains surrounding the blastocyst from which it "hatches" to commence implantation.

{{Fertilization Links}}
==Some Recent Findings==
[[File:Human_zygote_two_pronuclei_22.jpg|thumb|Human zygote inside the zone pellucida]]
[[File:Mouse_germinal_vesicle_04.jpg|thumb|alt=Mouse germinal vesicle|Mouse germinal vesicle{{#pmid:25144310|PMID25144310}}]]

{|
|-bgcolor="F5FAFF"
|

* '''Structure of Zona Pellucida Module Proteins'''{{#pmid:29853186|PMID29853186}} "The egg coat, an extracellular matrix made up of glycoprotein filaments, plays a key role in animal fertilization by acting as a gatekeeper for sperm. Egg coat components polymerize using a common zona pellucida (ZP) "domain" module that consists of two related immunoglobulin-like domains, called ZP-N and ZP-C. The ZP module has also been recognized in a large number of other secreted proteins with different biological functions, whose mutations are linked to severe human diseases. During the last decade, tremendous progress has been made toward understanding the atomic architecture of the ZP module and the structural basis of its polymerization. Moreover, sperm-binding regions at the N-terminus of mollusk and mammalian egg coat subunits were found to consist of domain repeats that also adopt a ZP-N fold. This discovery revealed an unexpected link between invertebrate and vertebrate fertilization and led to the first structure of an egg coat-sperm protein recognition complex. In this review we summarize these exciting findings, discuss their functional implications, and outline future challenges that must be addressed in order to develop a comprehensive view of this family of biomedically important extracellular molecules.

* '''Perivitelline threads in cleavage-stage human embryos: observations using time-lapse imaging'''{{#pmid:29074360|PMID29074360}} "Time-lapse imaging of the human preimplantation embryo in vitro has revealed a transient phenomenon involving the appearance of perivitelline threads, commonly observed at the two-cell stage. These threads span the perivitelline space, arising at the specific area where the cytoplasmic membrane contacts the zona pellucida, before any perivitelline space is formed. The threads persist as the cytoplasmic membrane retracts from the zona pellucida to form the first cleavage furrow. In this observational report, these structures and their incidence are described. ...Presence or absence of threads did not affect embryo development. This descriptive study is limited; further characterization of these structures is needed to elucidate their potential role in early human embryo development."

|}
{| class="wikitable mw-collapsible mw-collapsed"
! More recent papers  
|-
| [[File:Mark_Hill.jpg|90px|left]] {{Most_Recent_Refs}}

Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Zona+pellucida+development ''Zona pellucida Development''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Zona+pellucida ''Zona pellucida''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=zona+pellucida+birefringence ''zona pellucida birefringence''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=zona+pellucida+protein+1 ''zona pellucida protein 1''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=zona+pellucida+protein+2 ''zona pellucida protein 2''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=zona+pellucida+protein+3 ''zona pellucida protein 3'']

|}
{| class="wikitable mw-collapsible mw-collapsed"
! Older papers  
|-
| {{Older papers}}
* '''A single domain of the ZP2 zona pellucida protein mediates gamete recognition in mice and humans'''{{#pmid:21074836|PMID21074836}} "The extracellular zona pellucida surrounds ovulated eggs and mediates gamete recognition that is essential for mammalian fertilization. Zonae matrices contain three (mouse) or four (human) glycoproteins (ZP1-4), but which protein binds sperm remains controversial. ...These observations in transgenic mice document that the ZP2(51-149) sperm-binding domain is necessary for human and mouse gamete recognition and penetration through the zona pellucida."

* '''Biosynthesis of hamster zona pellucida is restricted to the oocyte'''{{#pmid:21074836|PMID21074836}} "In the present work, the expression of ZP1, ZP2, ZP3 and ZP4 is carefully analyzed by in situ hybridization (ISH) in hamster ovaries. Our data suggest that the four hamster ZP genes are expressed in a coordinate and oocyte-specific manner during folliculogenesis. Furthermore, this expression is maximal during the first stages of the oocyte development and declines in oocytes from later development stages, particularly within large antral follicles."

* '''Zona pellucida birefringence in in vivo and in vitro matured oocytes'''{{#pmid:20079896|PMID20079896}}

* '''Zona pellucida glycoprotein-1 binds to spermatozoa and induces acrosomal exocytosis'''{{#pmid:20504872|PMID20504872}} "These studies revealed for the first time that in humans ZP1, in addition to ZP3 and ZP4, binds to capacitated spermatozoa and induces acrosomal exocytosis."
|}
==Three-dimensional Structure==
[[File:Mouse- zona pellucida 01.jpg|thumb|Mouse zona pellucida{{#pmid:18539589|PMID18539589}}]]

Electron microscopic studies have shown a variety of appearances from a porous, net-like structure to a nearly smooth and compact structure.

An scanning electron microscopic review{{#pmid:16703610|PMID16703610}} of both human and mouse zona pellucida structures at ovulation show:

* spongy ZP appearance well correlates with mature oocytes.
* a delicate meshwork of thin interconnected filaments
* a regular alternating pattern of wide and tight meshes.
* wide meshes correspond to "pores" of the "spongy" ZP.
** tight meshes correspond to the compact parts of the ZP surrounding the pores.

==Human Zona Pellucida==

An immunochemistry study{{#pmid:18033806|PMID18033806}} of human ZP proteins expression during folliculogenesis within the ovary primordial follicle identified the presence of ZP1 and ZP3 in most follicles (93% and 95%, respectively), with ZP2 only in 32% of these follicles. The ZP proteins were also detected in the cytoplasm of cuboidal granulosa cells. This localization suggested to the authors that these proteins had been present since oogenesis.

[[File:Human embryo day 5 label.gif]]

==Mouse Zona Pellucida==

[[File:Mouse- zona pellucida 03.jpg|Mouse zona pellucida|600px]]

Mouse zona pellucida development.{{#pmid:18539589|PMID18539589}}

In the mouse ovarian oocytes, and not granulosa cells, are the only source for mouse ZP glycoproteins.<{{#pmid:11906903|PMID11906903}}

<gallery>
File:Mouse germinal vesicle 01.jpg|Oocyte and zona pellucida
File:Mouse germinal vesicle 02.jpg|Oocyte and granulosa cells
File:Mouse germinal vesicle 03.jpg|Oocyte and granulosa cells
File:Mouse germinal vesicle 04.jpg|Oocyte and granulosa cells
</gallery>
==Zona Pellucida Glycoprotein Features==
[[File:Human_oocyte_11.jpg|thumb|alt=Human oocyte contained inside zona pellucida|Human oocyte contained inside zona pellucida.{{#pmid:24586757|PMID24586757}}]]
# A "ZP domain", which is a signature domain comprised of approximately 260 amino acid (aa) residues.
# An N-terminal hydrophobic signal peptide sequence.
# A potential N- and O-linked glycosylation sites.
# A C-terminal hydrophobic transmembrane-like domain (TMD)
# A potential consensus proprotein convertase (furin) cleavage site (CFCS) upstream of transmembrane-like domain (TMD).

==Zona Pellucida glycoprotein 1==

(ZP1) A 638 amino acid glycoprotein located in the zona pellucida, synthesized by and surrounding the oocyte.

:'''Links:''' [http://www.omim.org/entry/195000 OMIM - ZP1]

==Zona Pellucida glycoprotein 2==
[[File:Model of gamete recognition zona pellucida.jpg|thumb|alt=Model of gamete recognition zona pellucida|Model of gamete recognition zona pellucida{{#pmid:24934154|PMID24934154}}]]
(ZP2) A 745 amino acid glycoprotein located in the {{zona pellucida}}, synthesized by and surrounding the {{oocyte}}. The protein acts as a secondary sperm receptor that binds sperm only after the induction of the sperm [[A#acrosome reaction|acrosome reaction]]. Before {{fertilization}} ZP2 binds spermatozoa. After {{fertilization}} ZP2 is proteolytically cleaved as an initial block to polyspermy.

:'''Links:''' [http://www.omim.org/entry/182888 OMIM - ZP2]

==Zona Pellucida glycoprotein 3==

{{Zona Pellucida Glycoprotein 3}} ({{ZP3}}) A 424 amino acid glycoprotein located in the {{zona pellucida}}, synthesized by and surrounding the {{oocyte}} required for initial zona matrix formation and during {{fertilization}} for species-specific sperm binding. Now thought to exist in 2 isoforms ZP3A and ZP3B (a second polymorphic allele).

A study using the {{mouse}} {{oocyte}},{{#pmid:28145526|PMID28145526}} has shown that ZP3 is also required for germinal vesicle breakdown in oocyte meiosis. ZP3 was initially expressed in the nucleus during prophase, and then translocates to the {{zona pellucida}}. Several potential ZP3-interacting proteins for the germinal vesicle effect were identified in this study: Protein tyrosine phosphatase, receptor type K (Ptprk), Aryl hydrocarbon receptor-interacting protein-like 1 (Aipl1), and Diaphanous related formin 2 (Diaph2).

:'''Links:''' [http://www.omim.org/entry/182889 OMIM - ZP3]

==Zona Pellucida glycoprotein 4==

(ZP4) A 540 amino acid glycoprotein located in the zona pellucida, synthesized by and surrounding the oocyte required for initial zona matrix formation and along with ZP3 during fertilization for inducing the acrosome reaction and inhibited the binding of spermatozoa to zona pellucida in a time- and dose-dependent reaction. Associated with several mammalian species (human, {{rat}}, hamster and {{rabbit}}) but not all mammalian species, the initial protein produced contains a N-terminal signal peptide sequence, a conserved ZP domain, a consensus furin cleavage site, and a C-terminal transmembrane domain. The furin cleavage may result in release of the mature protein from the plasma membrane for subsequent incorporation into the zona pellucida matrix.

:'''Links:''' [http://www.omim.org/entry/613514 OMIM - ZP4]

==Zona Pellucida Binding Protein==

(ZPBP) Not a zona pellucida protein, but a spermatozoa protein found located on the acrosome surface and one of several proteins that participate in secondary binding between acrosome-reacted sperm and the zona pellucida.

:'''Links:''' [http://www.omim.org/entry/608498 OMIM - ZPBP]

==Zona Pellucida Birefringence==

(ZPB) Optical property of the zona pellucida using polarization imaging when viewed microscopically. This property has been used to qualitatively predict the developmental potential of a in vitro matured metaphase-II (MII) oocytes.

High birefringence, compared to low, has been associated with oocytes contributing to conception cycles when compared with those of nonconception cycles and higher implantation, pregnancy, and live birth rates from transferred oocytes.{{#pmid:18284880|PMID18284880}}{{#pmid:20079896|PMID20079896}} This work follows on from the earlier discovery of an increased birefringence in the meiotic spindle an early indicator of oocyte activation{{#pmid:10859266|PMID10859266}}

'''Oocyte zona birefringence intensity is associated with embryonic implantation potential in ICSI cycles'''<ref name="PMID18284880" />

:"Using zona birefringence as the only selection criterion, two fertilized oocytes, preferably derived from high birefringence (HZB) oocytes, were selected for further culture and transfer. The required criteria were met by 135 ICSI cycles (124 patients; 34.9 ± 4.1 years of age). Embryos for transfer were used in 20 cycles derived from HZB/HZB oocytes, in 50 cycles from HZB/LZB oocytes and in 65 from LZB/LZB oocytes. The corresponding implantation (P < 0.025), pregnancy (P < 0.005) and live birth (P < 0.025) rates were significantly different between HZB/HZB and HZB/LZB versus LZB/LZB group. Embryo development was superior in embryos derived from HZB oocytes."

'''Zona pellucida birefringence in in vivo and in vitro matured oocytes'''<ref name="PMID20079896" />

The percentage of high birefringence oocytes was higher in immature than in mature oocytes (40.1 vs. 23.6%). Among immature oocytes, an increased percentage of HB in prophase-I stage oocytes compared to metaphase I stage oocytes was also observed (50.7 vs. 25.0%). However, the percentage of HB oocytes did not change when comparing oocytes before and after in vitro maturation for both prophase I and metaphase I oocytes. No influence of ZPB was observed on the spontaneous in vitro maturation potential. Exclusively for metaphase II retrieved oocytes, a positive influence of ZPB on fertilization (odds ratio [OR], 1.78; 95% confidence interval [CI], 1.27-2.49) and embryo quality (OR, 2.28; 95% CI, 1.04-4.99) was noted.

'''Birefringence''' - Defined as the double refraction of light in a transparent, molecularly ordered material, which is manifested by the existence of orientation-dependent differences in refractive index.

:'''Links:''' [http://www.olympusmicro.com/primer/lightandcolor/birefringence.html Olympus - Optical Birefringence]

==Abnormalities==

===Empty Follicle Syndrome===

Empty Follicle Syndrome (EFS) is clinically defined as the failure to aspirate oocytes from mature ovarian follicles during in vitro fertilization. At least one genetic cause has been identified, a missense mutation in {{zona pellucida}} protein 3 (ZP3).{{#pmid:28886344|PMID28886344}}

===Zona Pellucida Antibodies===
The zona pellucida has been shown to contain tissue-specific and species-specific antigens that can generate antibodies that inhibit sperm attachment. Anti-zona pellucida antibodies have been shown to occur in human serum though the cause of their generation is largely unknown.{{#pmid:2697213|PMID2697213}} It has been suggested that this may be a contributing factor to infertility{{#pmid:16372044|PMID16372044}} and premature ovarian failure.{{#pmid:17349637|PMID17349637}}

Note the generation of zona pellucida antibodies from specific antigens has also been used in controlling animal fertility.{{#pmid:16113040|PMID16113040}} This has also been suggested as a possible target in generating a contraceptive vaccine for controlling human fertility.{{#pmid:16113040|PMID16113040}}

==Terms==
* '''acrosome reaction''' - The chemical change within the {{spermatozoa}} following binding to the zona pellucida, that leads to the release of acrosomal enzymatic contents. These enzymes degrade the zona pellucida and allow a {{spermatozoa}} to penetrate an oocyte.

* '''granulosa cell''' - A specific cell type that proliferates in association with the oocyte within the developing [[F#follicle|follicles]] of the {{ovary}}. These cells form the [[F#follicle|follicle]] stratum granulosa and are also given specific names based upon their position within the [[F#follicle|follicle]]. In the [[A#antral_follicle|antral follicle]], [[M#membrana_granulosa|membrana granulosa]] sits on the [[F#follicular_basal_lamina|follicular basal lamina]] and lines the antrum as a stratified epithelium. The [[C#cumulus_oophorus|cumulus oophorus]] is a column of granulosa cells that attaches the oocyte to the [[F#follicle|follicle]] wall. The [[C#corona_radiata|corona radiata]] are the [[G#granulosa cell|granulosa cells]] that directly surround the oocyte, and are released along with it at ovulation. Following ovulation the corona radiata provide physical protection to the oocyte and granulosa cells within the ovulating [[F#follicle|follicle]] contribute to {{corpus luteum}}.

==References==

<references/>

===Reviews===
{{#pmid:29853181}}

{{#pmid:29699147}}

{{#pmid:25445843}}

{{#pmid:18539589}}

===Articles===

{{#pmid:29462288}}

{{#pmid:29094202}}

{{#pmid:28646452}}

{{#pmid:28886344}}

===Bookshelf===

* MBoC [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=mboc4.figgrp.3722 Figure 20-21. The zona pellucida]

'''Search NCBI Bookshelf''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=zona+pellucida zona pellucida] | [http://www.ncbi.nlm.nih.gov/sites/entrez?db=Books&cmd=search&term=acrosome+reaction acrosome reaction]

===Search Pubmed===

July 2010 "zona pellucida" All (4801) Review (582) Free Full Text (1408)

'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=zona%20pellucida zona pellucida] | [http://www.ncbi.nlm.nih.gov/sites/gquery?itool=toolbar&cmd=search&term=acrosome%20reaction acrosome reaction] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Zona+pellucida+protein+1 Zona pellucida protein 1] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Zona+pellucida+protein+2 Zona pellucida protein 2] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Zona+pellucida+protein+3 Zona pellucida protein 3] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Zona+pellucida+protein+4 Zona pellucida protein 4]

'''Search Images:''' [http://www.ncbi.nlm.nih.gov/pmc/?term=zona+pellucida&report=imagesdocsum zona pellucida]

==Additional Images==
<gallery>
File:Mouse-_zona_pellucida_01.jpg|[[Mouse Development|Mouse]] oocyte and zona pellucida - light and TEM
File:Mouse_oocyte_and_zona_pellucida_EM01a.jpg|[[Mouse Development|Mouse]] oocyte and zona pellucida TEM
File:Cat_oocyte_zona_pellucida_01.jpg|[[Cat Development|Cat]] oocyte zona pellucida SEM
File:Cat_oocyte_zona_pellucida_02.jpg|[[Cat Development|Cat]] oocyte zona pellucida SEM
File:Cat_spermatozoa_bound_to_oocyte_zona_pellucida.jpg|[[Cat Development|Cat]] oocyte zona pellucida spermatozoa bound SEM
File:Hamster_oocyte_zona_pellucida_SEM.jpg|[[Hamster_Development|Hamster]] oocyte zona pellucida SEM
File:Patterns_of_ZPC_Deposition_in_Porcine_Oocytes.jpg|[[Pig Development|Pig]] ZPC deposition in oocyte-cumulus complexes
File:Bailey001.jpg|Historic - From a section of the ovary of a 12-year old girl
</gallery>

==External Links==
{{External Links}}

* OMIM [http://www.omim.org/entry/195000 ZP1] | [http://www.omim.org/entry/182888 ZP2] | [http://www.omim.org/entry/182889 ZP3] | [http://www.omim.org/entry/613514 ZP4] | [http://www.omim.org/entry/608498 ZPBP]

{{Glossary}}

{{Footer}}
[[Category:Week 1]] [[Category:Zona Pellucida]] [[Category:Oocyte]]

Template talk:ZP1 - Revision history

Z8600021: Blanked the page

Z8600021: Undo revision 376941 by Z8600021 (talk)

Z8600021: Blanked the page

Z8600021 at 10:59, 15 April 2019

Z8600021: Created page with " {{Header}} ==Introduction== Oocyte and developing zona pellucida in the ovary I..."