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Cell membrane
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Illustration of a Eukaryotic cell membrane
Comparison of Eukaryotes vs. Prokaryotes
The cell membrane (also known as the plasma membrane or cytoplasmic membrane) is a biological membrane that separates the interior of all cells from the outside environment.[1][2] The cell membrane is selectively permeable to ions and organic molecules and controls the movement of substances in and out of cells.[3] The basic function of the cell membrane is to protect the cell from its surroundings.
It consists of the lipid bilayer with embedded proteins. Cell membranes are involved in a variety of cellular processes such as cell adhesion, ion conductivity and cell signalling and serve as the attachment surface for several extracellular structures, including the cell wall, glycocalyx, and intracellular cytoskeleton. Cell membranes can be artificially reassembled.[4][5][6]
Contents
[hide]
1 History
2 Function
3 Prokaryotes
4 Structures
4.1 Fluid mosaic model
4.2 Lipid bilayer
4.3 Membrane polarity
4.4 Membrane structures
4.5 Cytoskeleton
5 Composition
5.1 Lipids
5.2 Phospholipids forming lipid vesicles
5.3 Carbohydrates
5.4 Proteins
6 Variation
7 Permeability
8 See also
9 Notes and references
10 External links
History
Main article: History of cell membrane theory
The structure has been variously referred to by different writers as the ectoplast (de Vries, 1885),[7] Plasmahaut (plasma skin, Pfeffer, 1877, 1891),[8] Hautschicht (skin layer, Pfeffer, 1886; used with a different meaning by Hofmeister, 1867), plasmatic membrane (Pfeffer, 1900),[9] plasma membrane, cytoplasmic membrane, cell envelope and cell membrane.[10][11]
Some authors that did not believe that there was a functional permeable boundary at the surface of the cell preferred to use the term plasmalemma (coined by Mast, 1924) for the external region of the cell.[12][13][14]
Function
A detailed diagram of the cell membrane
Illustration depicting cellular diffusion
The cell membrane (or plasma membrane or plasmalemma) surrounds the cytoplasm of living cells, physically separating the intracellular components from the extracellular environment. The cell membrane also plays a role in anchoring the cytoskeleton to provide shape to the cell, and in attaching to the extracellular matrix and other cells to hold them together to form tissues. Fungi, bacteria, most archaea, and plants also have a cell wall, which provides a mechanical support to the cell and precludes the passage of larger molecules.
The cell membrane is selectively permeable and able to regulate what enters and exits the cell, thus facilitating the transport of materials needed for survival. The movement of substances across the membrane can be either "passive", occurring without the input of cellular energy, or "active", requiring the cell to expend energy in transporting it. The membrane also maintains the cell potential. The cell membrane thus works as a selective filter that allows only certain things to come inside or go outside the cell. The cell employs a number of transport mechanisms that involve biological membranes:
1. Passive osmosis and diffusion: Some substances (small molecules, ions) such as carbon dioxide (CO2) and oxygen (O2), can move across the plasma membrane by diffusion, which is a passive transport process. Because the membrane acts as a barrier for certain molecules and ions, they can occur in different concentrations on the two sides of the membrane. Such a concentration gradient across a semipermeable membrane sets up an osmotic flow for the water.
2. Transmembrane protein channels and transporters: Nutrients, such as sugars or amino acids, must enter the cell, and certain products of metabolism must leave the cell. Such molecules diffuse passively through protein channels such as aquaporins (in the case of water (H2O)) in facilitated diffusion or are pumped across the membrane by transmembrane transporters. Protein channel proteins, also called permeases, are usually quite specific, recognizing and transporting only a limited food group of chemical substances, often even only a single substance.
3. Endocytosis: Endocytosis is the process in which cells absorb molecules by engulfing them. The plasma membrane creates a small deformation inward, called an invagination, in which the substance to be transported is captured. The deformation then pinches off from the membrane on the inside of the cell, creating a vesicle containing the captured substance. Endocytosis is a pathway for internalizing solid particles ("cell eating" or phagocytosis), small molecules and ions ("cell drinking" or pinocytosis), and macromolecules. Endocytosis requires energy and is thus a form of active transport.
4. Exocytosis: Just as material can be brought into the cell by invagination and formation of a vesicle, the membrane of a vesicle can be fused with the plasma membrane, extruding its contents to the surrounding medium. This is the process of exocytosis. Exocytosis occurs in various cells to remove undigested residues of substances brought in by endocytosis, to secrete substances such as hormones and enzymes, and to transport a substance completely across a cellular barrier. In the process of exocytosis, the undigested waste-containing food vacuole or the secretory vesicle budded from Golgi apparatus, is first moved by cytoskeleton from the interior of the cell to the surface. The vesicle membrane comes in contact with the plasma membrane. The lipid molecules of the two bilayers rearrange themselves and the two membranes are, thus, fused. A passage is formed in the fused membrane and the vesicles discharges its contents outside the cell.
Prokaryotes
Gram-negative bacteria have both a plasma membrane and an outer membrane separated by periplasm. Other prokaryotes have only a plasma membrane. Bacteria are also surrounded by a cell wall composed of peptidoglycan (amino acids and sugars). Some eukaryotic cells also have cell walls, but none that are made of peptidoglycan. The outer membrane of gram negative bacteria is rich in lipopolysaccharides. The outer membrane can bleb out into periplasmic protrusions under stress conditions or upon virulence requirements while encountering a host target cell, and thus such blebs may work as virulence organelles.[15]
Structures
Fluid mosaic model
According to the fluid mosaic model of S. J. Singer and G. L. Nicolson (1972), which replaced the earlier model of Davson and Danielli, biological membranes can be considered as a two-dimensional liquid in which lipid and protein molecules diffuse more or less easily.[16] Although the lipid bilayers that form the basis of the membranes do indeed form two-dimensional liquids by themselves, the plasma membrane also contains a large quantity of proteins, which provide more structure. Examples of such structures are protein-protein complexes, pickets and fences formed by the actin-based cytoskeleton, and potentially lipid rafts.
Lipid bilayer
Diagram of the arrangement of amphipathic lipid molecules to form a lipid bilayer. The yellow polar head groups separate the grey hydrophobic tails from the aqueous cytosolic and extracellular environments.
Lipid bilayers form through the process of self-assembly. The cell membrane consists primarily of a thin layer of amphipathic phospholipids that spontaneously arrange so that the hydrophobic "tail" regions are isolated from the surrounding water while the hydrophilic "head" regions interact with the intracellular (cytosolic) and extracellular faces of the resulting bilayer. This forms a continuous, spherical lipid bilayer. Hydrophobic interactions (also known as the hydrophobic effect) are the major driving forces in the formation of lipid bilayers. An increase in interactions between hydrophobic molecules (causing clustering of hydrophobic regions) allows water molecules to bond more freely with each other, increasing the entropy of the system. This complex interaction can include noncovalent interactions such as van der Waals, electrostatic and hydrogen bonds.
Lipid bilayers are generally impermeable to ions and polar molecules. The arrangement of hydrophilic heads and hydrophobic tails of the lipid bilayer prevent polar solutes (ex. amino acids, nucleic acids, carbohydrates, proteins, and ions) from diffusing across the membrane, but generally allows for the passive diffusion of hydrophobic molecules. This affords the cell the ability to control the movement of these substances via transmembrane protein complexes such as pores, channels and gates. Flippases and scramblases concentrate phosphatidyl serine, which carries a negative charge, on the inner membrane. Along with NANA, this creates an extra barrier to charged moieties moving through the membrane.
Membranes serve diverse functions in eukaryotic and prokaryotic cells. One important role is to regulate the movement of materials into and out of cells. The phospholipid bilayer structure (fluid mosaic model) with specific membrane proteins accounts for the selective permeability of the membrane and passive and active transport mechanisms. In addition, membranes in prokaryotes and in the mitochondria and chloroplasts of eukaryotes facilitate the synthesis of ATP through chemiosmosis.
Membrane polarity
See also: Epithelial polarity
Alpha intercalated cell
The apical membrane of a polarized cell is the surface of the plasma membrane that faces inward to the lumen. This is particularly evident in epithelial and endothelial cells, but also describes other polarized cells, such as neurons. The basolateral membrane of a polarized cell is the surface of the plasma membrane that forms its basal and lateral surfaces. It faces outwards, towards the interstitium, and away from the lumen. Basolateral membrane is a compound phrase referring to the terms "basal (base) membrane" and "lateral (side) membrane", which, especially in epithelial cells, are identical in composition and activity. Proteins (such as ion channels and pumps) are free to move from the basal to the lateral surface of the cell or vice versa in accordance with the fluid mosaic model. Tight junctions join epithelial cells near their apical surface to prevent the migration of proteins from the basolateral membrane to the apical membrane. The basal and lateral surfaces thus remain roughly equivalent[clarification needed] to one another, yet distinct from the apical surface.
Membrane structures
Diagram of the Cell Membrane's structures and their function.
Cell membrane can form different types of "supramembrane" structures such as caveola, postsynaptic density, podosome, invadopodium, focal adhesion, and different types of cell junctions. These structures are usually responsible for cell adhesion, communication, endocytosis and exocytosis. They can be visualized by electron microscopy or fluorescence microscopy. They are composed of specific proteins, such as integrins and cadherins.
Cytoskeleton
The cytoskeleton is found underlying the cell membrane in the cytoplasm and provides a scaffolding for membrane proteins to anchor to, as well as forming organelles that extend from the cell. Indeed, cytoskeletal elements interact extensively and intimately with the cell membrane.[17] Anchoring proteins restricts them to a particular cell surface — for example, the apical surface of epithelial cells that line the vertebrate gut — and limits how far they may diffuse within the bilayer. The cytoskeleton is able to form appendage-like organelles, such as cilia, which are microtubule-based extensions covered by the cell membrane, and filopodia, which are actin-based extensions. These extensions are ensheathed in membrane and project from the surface of the cell in order to sense the external environment and/or make contact with the substrate or other cells. The apical surfaces of epithelial cells are dense with actin-based finger-like projections known as microvilli, which increase cell surface area and thereby increase the absorption rate of nutrients. Localized decoupling of the cytoskeleton and cell membrane results in formation of a bleb.
Composition
Cell membranes contain a variety of biological molecules, notably lipids and proteins. Material is incorporated into the membrane, or deleted from it, by a variety of mechanisms:
Fusion of intracellular vesicles with the membrane (exocytosis) not only excretes the contents of the vesicle but also incorporates the vesicle membrane's components into the cell membrane. The membrane may form blebs around extracellular material that pinch off to become vesicles (endocytosis).
If a membrane is continuous with a tubular structure made of membrane material, then material from the tube can be drawn into the membrane continuously.
Although the concentration of membrane components in the aqueous phase is low (stable membrane components have low solubility in water), there is an exchange of molecules between the lipid and aqueous phases.
Lipids
Examples of the major membrane phospholipids and glycolipids: phosphatidylcholine (PtdCho), phosphatidylethanolamine (PtdEtn), phosphatidylinositol (PtdIns), phosphatidylserine (PtdSer).
The cell membrane consists of three classes of amphipathic lipids: phospholipids, glycolipids, and sterols. The amount of each depends upon the type of cell, but in the majority of cases phospholipids are the most abundant.[18] In RBC studies, 30% of the plasma membrane is lipid.
The fatty chains in phospholipids and glycolipids usually contain an even number of carbon atoms, typically between 16 and 20. The 16- and 18-carbon fatty acids are the most common. Fatty acids may be saturated or unsaturated, with the configuration of the double bonds nearly always "cis". The length and the degree of unsaturation of fatty acid chains have a profound effect on membrane fluidity[19] as unsaturated lipids create a kink, preventing the fatty acids from packing together as tightly, thus decreasing the melting temperature (increasing the fluidity) of the membrane. The ability of some organisms to regulate the fluidity of their cell membranes by altering lipid composition is called homeoviscous adaptation.
The entire membrane is held together via non-covalent interaction of hydrophobic tails, however the structure is quite fluid and not fixed rigidly in place. Under physiological conditions phospholipid molecules in the cell membrane are in the liquid crystalline state. It means the lipid molecules are free to diffuse and exhibit rapid lateral diffusion along the layer in which they are present. However, the exchange of phospholipid molecules between intracellular and extracellular leaflets of the bilayer is a very slow process. Lipid rafts and caveolae are examples of cholesterol-enriched microdomains in the cell membrane. Also, a fraction of the lipid in direct contact with integral membrane proteins, which is tightly bound to the protein surface is called annular lipid shell; it behaves as a part of protein complex.
In animal cells cholesterol is normally found dispersed in varying degrees throughout cell membranes, in the irregular spaces between the hydrophobic tails of the membrane lipids, where it confers a stiffening and strengthening effect on the membrane.[3]
Phospholipids forming lipid vesicles
Lipid vesicles or liposomes are circular pockets that are enclosed by a lipid bilayer. These structures are used in laboratories to study the effects of chemicals in cells by delivering these chemicals directly to the cell, as well as getting more insight into cell membrane permeability. Lipid vesicles and liposomes are formed by first suspending a lipid in an aqueous solution then agitating the mixture through sonication, resulting in a vesicle. By measuring the rate of efflux from that of the inside of the vesicle to the ambient solution, allows researcher to better understand membrane permeability. Vesicles can be formed with molecules and ions inside the vesicle by forming the vesicle with the desired molecule or ion present in the solution. Proteins can also be embedded into the membrane through solubilizing the desired proteins in the presence of detergents and attaching them to the phospholipids in which the liposome is formed. These provide researchers with a tool to examine various membrane protein functions.
Carbohydrates
Plasma membranes also contain carbohydrates, predominantly glycoproteins, but with some glycolipids (cerebrosides and gangliosides). For the most part, no glycosylation occurs on membranes within the cell; rather generally glycosylation occurs on the extracellular surface of the plasma membrane. The glycocalyx is an important feature in all cells, especially epithelia with microvilli. Recent data suggest the glycocalyx participates in cell adhesion, lymphocyte homing, and many others. The penultimate sugar is galactose and the terminal sugar is sialic acid, as the sugar backbone is modified in the Golgi apparatus. Sialic acid carries a negative charge, providing an external barrier to charged particles.
Proteins
Type Description Examples
Integral proteins
or transmembrane proteins Span the membrane and have a hydrophilic cytosolic domain, which interacts with internal molecules, a hydrophobic membrane-spanning domain that anchors it within the cell membrane, and a hydrophilic extracellular domain that interacts with external molecules. The hydrophobic domain consists of one, multiple, or a combination of α-helices and β sheet protein motifs. Ion channels, proton pumps, G protein-coupled receptor
Lipid anchored proteins Covalently bound to single or multiple lipid molecules; hydrophobically insert into the cell membrane and anchor the protein. The protein itself is not in contact with the membrane. G proteins
Peripheral proteins Attached to integral membrane proteins, or associated with peripheral regions of the lipid bilayer. These proteins tend to have only temporary interactions with biological membranes, and once reacted, the molecule dissociates to carry on its work in the cytoplasm. Some enzymes, some hormones
The cell membrane has large content of proteins, typically around 50% of membrane volume[19] These proteins are important for cell because they are responsible for various biological activities. Approximately a third of the genes in yeast code specifically for them, and this number is even higher in multicellular organisms.[18]
The cell membrane, being exposed to the outside environment, is an important site of cell–cell communication. As such, a large variety of protein receptors and identification proteins, such as antigens, are present on the surface of the membrane. Functions of membrane proteins can also include cell–cell contact, surface recognition, cytoskeleton contact, signaling, enzymatic activity, or transporting substances across the membrane.
Most membrane proteins must be inserted in some way into the membrane. For this to occur, an N-terminus "signal sequence" of amino acids directs proteins to the endoplasmic reticulum, which inserts the proteins into a lipid bilayer. Once inserted, the proteins are then transported to their final destination in vesicles, where the vesicle fuses with the target membrane.
Variation
The cell membrane has different lipid and protein compositions in distinct types of cells and may have therefore specific names for certain cell types:
Sarcolemma in myocytes
Oolemma in oocytes
Axolemma in neuronal processes - axons
Historically, the plasma membrane was also referred to as the plasmalemma
Permeability
The permeability of a membrane is the rate of passive diffusion of molecules through the membrane. These molecules are known as permeant molecules. Permeability depends mainly on the electric charge and polarity of the molecule and to a lesser extent the molar mass of the molecule. Due to the cell membrane's hydrophobic nature, small electrically neutral molecules pass through the membrane more easily than charged, large ones. The inability of charged molecules to pass through the cell membrane results in pH partition of substances throughout the fluid compartments of the body.
See also
Ammonium transporter
Annular lipid shell
AP2 adaptors
Artificial cell
Bacterial cell structure
Bangstad syndrome
Cell cortex
Cell damage, including damage to cell membrane
Cell theory
Cytoneme
Elasticity of cell membranes
Gram-positive bacteria
Membrane models
Membrane nanotubule
History of cell membrane theory
Lipid raft
Trogocytosis
Notes and references
Jump up ^ Kimball's Biology pages, Cell Membranes
Jump up ^ Singleton P (1999). Bacteria in Biology, Biotechnology and Medicine (5th ed.). New York: Wiley. ISBN 0-471-98880-4.
^ Jump up to: a b Alberts B, Johnson A, Lewis J, et al. (2002). Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN 0-8153-3218-1.
Jump up ^ Budin, Itay; Devaraj, Neal K. (December 29, 2011). "Membrane Assembly Driven by a Biomimetic Coupling Reaction". Journal of the American Chemical Society. 134 (2): 751–753. doi:10.1021/ja2076873. Retrieved February 18, 2012.
Jump up ^ Staff (January 25, 2012). "Chemists Synthesize Artificial Cell Membrane". ScienceDaily. Retrieved February 18, 2012.
Jump up ^ Staff (January 26, 2012). "Chemists create artificial cell membrane". kurzweilai.net. Retrieved February 18, 2012.
Jump up ^ de Vries, H. 1885. Plasmolytische Studien über die Wand der Vakuolen. Jahrb. wiss. Bot. 16, 465–598
Jump up ^ Pfeffer, W. 1877. Osmotische Untersuchungen: Studien zur Zell Mechanik. Engelmann, Leipzig.
Jump up ^ Pfeffer, W., 1900–1906. The Physiology of Plants, [1]. Translated by A. J. Ewart from the 2nd German ed. of Pflanzenphysiologie, 1897-1904, [2]. Clarendon Press, Oxford.
Jump up ^ Sharp, L. W. (1921). Introduction To Cytology. New York: McGraw Hill, p. 42.
Jump up ^ Kleinzeller, A. 1999. Charles Ernest Overton’s concept of a cell membrane. In: Membrane permeability: 100 years since Ernest Overton (ed. Deamer D.W., Kleinzeller A., Fambrough D.M.), pp. 1–18, Academic Press, San Diego, [3].
Jump up ^ Mast, S. O. 1924. Structure and locomotion in Amoeba proteus. Anat. Rec. 29: 88, [4].
Jump up ^ Plowe, J. Q. (1931). Membranes in the plant cell. I. Morphological membranes at protoplasmic surfaces. Protoplasma 12, 196-220, [5].
Jump up ^ Wayne, R. 2009. Plant Cell Biology: From Astronomy to Zoology. Amsterdam: Elsevier/Academic Press, p. 17.
Jump up ^ YashRoy, R.C. (1999). "A structural model for virulence organellae of gram negative organisms with reference to Salmonella pathogenicity in chicken ileum". Indian Journal of Poultry Science. 34 (2): 213–219.
Jump up ^ Singer SJ, Nicolson GL (Feb 1972). "The fluid mosaic model of the structure of cell membranes". Science. 175 (4023): 720–31. PMID 4333397. doi:10.1126/science.175.4023.720.
Jump up ^ Doherty GJ, McMahon HT (2008). "Mediation, Modulation and Consequences of Membrane-Cytoskeleton Interactions". Annual Review of Biophysics. 37: 65–95. PMID 18573073. doi:10.1146/annurev.biophys.37.032807.125912.
^ Jump up to: a b Lodish H, Berk A, Zipursky LS, et al. (2000). Molecular Cell Biology (4th ed.). New York: Scientific American Books. ISBN 0-7167-3136-3.
^ Jump up to: a b Jesse Gray; Shana Groeschler; Tony Le; Zara Gonzalez (2002). "Membrane Structure" (SWF). Davidson College. Retrieved 2007-01-11.
External links
Wikimedia Commons has media related to Cell membrane.
Lipids, Membranes and Vesicle Trafficking - The Virtual Library of Biochemistry and Cell Biology
Cell membrane protein extraction protocol
Membrane homeostasis, tension regulation, mechanosensitive membrane exchange and membrane traffic
3D structures of proteins associated with plasma membrane of eukaryotic cells
Lipid composition and proteins of some eukariotic membranes
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Structures of the cell / organelles (TH H1.00.01.2-3)
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NDL: 00569973
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2018 ACC Men's Basketball Tournament
From Wikipedia, the free encyclopedia
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2018 ACC Men's Basketball Tournament
2018 ACC Tournament Logo
Classification
Division I
Season
2017–18
Teams
15
Site
Barclays Center
Brooklyn, New York
Champions
Virginia (3rd title)
Winning coach
Tony Bennett (2nd title)
MVP
Kyle Guy (Virginia)
Attendance
82,390
Television
ESPN, ESPN2, ESPNU, ACCN
ACC Men's Basketball Tournaments
← 2017
2019 →
2017–18 ACC men's basketball standings
v
t
e
Conf
Overall
Team
W
L
PCT
W
L
PCT
No. 1 Virginia † 17 – 1 .944 31 – 3 .912
No. 9 Duke 13 – 5 .722 29 – 8 .784
No. 22 Miami (FL) 11 – 7 .611 22 – 10 .688
No. 20 Clemson 11 – 7 .611 25 – 10 .714
NC State 11 – 7 .611 21 – 12 .636
No. 10 North Carolina 11 – 7 .611 26 – 11 .703
Virginia Tech 10 – 8 .556 21 – 12 .636
Florida State 9 – 9 .500 23 – 12 .657
Louisville 9 – 9 .500 22 – 14 .611
Notre Dame 8 – 10 .444 21 – 15 .583
Syracuse 8 – 10 .444 23 – 14 .622
Boston College 7 – 11 .389 19 – 16 .543
Georgia Tech 6 – 12 .333 13 – 19 .406
Wake Forest 4 – 14 .222 11 – 20 .355
Pittsburgh 0 – 18 .000 8 – 24 .250
† 2018 ACC Tournament winner
Rankings from AP Poll
The 2018 ACC Men's Basketball Tournament was the postseason men's basketball tournament for the Atlantic Coast Conference held at Barclays Center in Brooklyn, New York from March 6–10, 2018. It was the 65th annual edition of the tournament, and the second year in a row being held at Barclays Center. The Virginia Cavaliers entered the tournament as the top seed, with a 17–1 conference record (28–2 overall) under the guidance of Tony Bennett. UVA also began the tournament unanimously ranked No. 1 in the country in both major polls.
The Cavaliers went on to win the tournament by handily defeating Louisville 75–58, Clemson 64–58, and North Carolina 71–63 in the championship game. Sophomore guard Kyle Guy was named Tournament MVP. Games were shown on over-the-air television in local media markets by the syndicated ACC Network and simulcast nationally on various ESPN cable networks.
Both the Virginia–North Carolina title game and Duke–North Carolina semifinal game set the Barclays Center attendance record for college basketball games, and conference leadership vowed to return the ACC Tournament to New York in the near future.[1][2]
Contents
[hide]
1 Seeds
2 Schedule
3 Bracket
4 Game summaries
4.1 First round
4.2 Second round
4.3 Quarterfinals
4.4 Semifinals
4.5 Championship
5 Awards and honors
6 See also
7 References
Seeds[edit]
All 15 ACC teams participated in the tournament. Teams were seeded by record within the conference, with a tiebreaker system to seed teams with identical conference records. The tournament was held over five consecutive days. Teams ranked 10–15 played on the first day; teams ranked 5–9 entered the bracket on the second day, with teams ranked 1–4 entering on the third day (quarterfinals). The semifinals were played on the fourth day, and the finals on the last day.[3]
Seed
School
Conference
Record
Tiebreaker
1 Virginia 17–1
2 Duke 13–5
3 Miami 11–7 2–1 vs Clem/NCST/UNC
4 Clemson 11–7 3–2 vs Miami/NCST/UNC
5 NC State 11–7 1–0 vs Duke
6 North Carolina 11–7 1–1 vs Duke
7 Virginia Tech 10–8
8 Florida State 9–9 3–3 vs Miami/Clem/NCST/UNC
9 Louisville 9–9 0–4 vs Miami/Clem/NCST/UNC
10 Notre Dame 8–10 1–0 vs Syracuse
11 Syracuse 8–10 0–1 vs ND
12 Boston College 7–11
13 Georgia Tech 6–12
14 Wake Forest 4–14
15 Pittsburgh 0–18
Schedule[edit]
All games will be televised on the ACC Network within the ACC footprint and simulcast nationally on the ESPN networks denoted below.
Session
Game
Time
Matchup
Score
Television
Attendance
First round – Tuesday, March 6
Opening day 1 12:00 pm No. 12 Boston College vs No. 13 Georgia Tech 87–77 ESPN2 10,612[4]
2 2:00 pm No. 10 Notre Dame vs No. 15 Pittsburgh 67–64
3 7:00 pm No. 11 Syracuse vs No. 14 Wake Forest 73–64 ESPNU
Second round – Wednesday, March 7
1 4 12:00 pm No. 8 Florida State vs No. 9 Louisville 74–82 ESPN 17,732[5]
5 2:30 pm No. 5 NC State vs No. 12 Boston College 87–91
2 6 7:00 pm No. 7 Virginia Tech vs No. 10 Notre Dame 65–71 ESPN2
7 9:30 pm No. 6 North Carolina vs No. 11 Syracuse 78–59
Quarterfinals – Thursday, March 8
3 8 12:00 pm No. 1 Virginia vs No. 9 Louisville 75–58 ESPN 17,732[6]
9 2:00 pm No. 4 Clemson vs No. 12 Boston College 90–82
4 10 7:00 pm No. 2 Duke vs No. 10 Notre Dame 88–70
11 9:00 pm No. 3 Miami vs No. 6 North Carolina 65–82
Semifinals – Friday, March 9
5 12 7:00 pm No. 1 Virginia vs No. 4 Clemson 64–58 ESPN2 18,157[7]
13 9:00 pm No. 2 Duke vs No. 6 North Carolina 69–74 ESPN
Championship – Saturday, March 10
6 14 8:30 pm No. 1 Virginia vs No. 6 North Carolina 71–63 ESPN 18,157[8]
Game times in ET. Rankings denote tournament seed.
Bracket[edit]
[9]
First round
Tuesday, March 6
ESPN2/ESPNU/ACCN Second round
Wednesday, March 7
ESPN/ESPN2/ACCN Quarterfinals
Thursday, March 8
ESPN/ESPN2/ACCN Semifinals
Friday, March 9
ESPN/ESPN2/ACCN Championship
Saturday, March 10
ESPN/ACCN
1 Virginia 75
9 Louisville 58
8 Florida State 74
9 Louisville 82
1 Virginia 64
4 Clemson 58
4 Clemson 90
12 Boston College 82
5 NC State 87
12 Boston College 91
12 Boston College 87
13 Georgia Tech 77
1 Virginia 71
6 North Carolina 63
2 Duke 88
10 Notre Dame 70
7 Virginia Tech 65
10 Notre Dame 71
10 Notre Dame 67
15 Pittsburgh 64
2 Duke 69
6 North Carolina 74
3 Miami 65
6 North Carolina 82
6 North Carolina 78
11 Syracuse 59
11 Syracuse 73
14 Wake Forest 64
Game summaries[edit]
First round[edit]
Three games were played in the first round. In all three, the higher seed advanced.
In the first game, the 12-seed Boston College Eagles took on the 13-seed Georgia Tech Yellow Jackets. Boston College got out to ten point lead at half time. They opened the second half on a 15-5 run, bring their lead to 20 with under 14 minutes to play. A late flurry of scoring by Georgia Tech reduced the Eagles lead to 6 with only 45 seconds remaining, but BC hit four free throws at the end of the game, ending the chance of a Yellow Jacket comeback. Georgia Tech's Tadric Jackson led all scorers with 29 points in the losing effort.[10]
The second game featured the 10-seed Notre Dame Fighting Irish against the 15-seed Pittsburgh Panthers. The game looked to be one of the biggest mismatches in the tournament, with Notre Dame likely post-season bound with a 20-win season and Pittsburgh entering the tournament having lost every conference game and sporting a woeful 8-24 record. Instead, the two teams played the closest game of the first round, with Notre Dame clinging to a two-point lead with less than a minute remaining; however a controversial loose-ball foul against Pitt's Marcus Carr followed by a turnover by Carr allowed Notre Dame to preserve their lead and close out the game 67-64. Carr's late game mistakes were ironic as his three-pointer was the one that cut the Notre Dame lead to two, and he scored 18 in the game, tied with Jared Wilson-Frame to lead the Panthers in scoring. Notre Dame's Bonzie Colson led all scorers with 19.[11]
The last game of the day featured the 11-seed Syracuse Orange versus the 14-seed Wake Forest Demon Deacons. Syracuse was playing with a short bench, as several injuries limited their ability to substitute. Despite leading by 19 with under eight minutes to play, the wear on Syracuse's starters showed, as they played all but the last 32 seconds of the second half, and that substitution was forced by Tyus Battle's fifth foul of the game. The Demon Deacons had cut the lead to six with fresher legs, but the Orange held on to win 73-64. Wake Forest's Bryant Crawford led all scoring with 22 points.[12]
ESPN2
March 6
12:00 pm
Report
No. 12 Boston College 87, No. 13 Georgia Tech 77
Scoring by half: 36–26, 51–51
Pts: Bowman (26)
Rebs: Mitchell (11)
Asts: Bowman (6) Pts: Jackson (29)
Rebs: Gueye (8)
Asts: Jackson (5)
Barclays Center
Brooklyn, NY
Attendance: 10,612
Referees: Bert Smith, Les Jones, AJ Dasai
ESPN2
March 6
2:00 pm
Report
No. 10 Notre Dame 67, No. 15 Pittsburgh 64
Scoring by half: 36–24, 31–40
Pts: Colson (19)
Rebs: Geben, Mooney (9)
Asts: Farrell (3) Pts: Carr, Wilson-Frame (18)
Rebs: Stewart (5)
Asts: Stevenson (4)
Barclays Center
Brooklyn, NY
Attendance: 10,612
Referees: Lee Cassell, Jerry Heater, Clarence Armstrong
ESPNU
March 6
7:00 pm
Report
No. 11 Syracuse 73, No. 14 Wake Forest 64
Scoring by half: 30–24, 43–40
Pts: Dolezaj (20)
Rebs: Brissett, Chukwu (9)
Asts: Howard (5) Pts: Crawford (22)
Rebs: Okeke (6)
Asts: Crawford (6)
Barclays Center
Brooklyn, NY
Attendance: 10,612
Referees: Bill Covington Jr., Raymond Styons, Tim Comer
Second round[edit]
ESPN
March 7
12:00 pm
Report
No. 8 Florida State 74, No. 9 Louisville 82
Scoring by half: 22–41, 52–41
Pts: Forrest, Savoy (14)
Rebs: Kabengele, Mann (6)
Asts: Forrest (6) Pts: Snider (19)
Rebs: Adel (8)
Asts: Snider (6)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Mike Eades, Tim Nestor, AJ Desai
ESPN
March 7
2:00 pm
Report
No. 5 NC State 87, No. 12 Boston College 91
Scoring by half: 31–45, 56–46
Pts: Freeman (21)
Rebs: Yurtseven (9)
Asts: Johnson (6) Pts: Robinson (26)
Rebs: Mitchell (15)
Asts: Robinson (5)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Jamie Luckie, Ron Groover, Jerry Heater
ESPN2
March 7
7:00 pm
Report
No. 7 Virginia Tech 65, No. 10 Notre Dame 71
Scoring by half: 34–21, 31–50
Pts: Robinson, Bibbs (15)
Rebs: Blackshear Jr. (8)
Asts: Robinson (7) Pts: Farrell (22)
Rebs: Pfleuger (9)
Asts: Gibbs (7)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Ted Valentine, Raymond Styons, Les Jones
ESPN2
March 7
9:00 pm
Report
No. 6 North Carolina 78, No. 11 Syracuse 59
Scoring by half: 39–29, 39–30
Pts: Williams (17)
Rebs: Pinson (11)
Asts: Pinson (6) Pts: Brissett (20)
Rebs: Brissett (10)
Asts: Howard (4)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Roger Ayers, Bert Smith, Clarence Armstrong
Quarterfinals[edit]
ESPN
March 8
12:00 pm
Report
No. 1 Virginia 75, No. 9 Louisville 58
Scoring by half: 38–27, 37–32
Pts: Guy (19)
Rebs: Guy (7)
Asts: Hall (5) Pts: Spalding (16)
Rebs: Spalding (6)
Asts: Spalding, Snider (3)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Ted Valentine, Jamie Luckie, Tim Comer
ESPN
March 8
2:00 pm
Report
No. 4 Clemson 90, No. 12 Boston College 82
Scoring by half: 43–36, 47–46
Pts: DeVoe (25)
Rebs: Thomas (12)
Asts: DeVoe, Reed (4) Pts: Bowman (23)
Rebs: Mitchell (9)
Asts: Bowman (4)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Mike Eades, Tim Nestor, Les Jones
ESPN
March 8
7:00 pm
Report
No. 2 Duke 88, No. 10 Notre Dame 70
Scoring by half: 41–37, 47–33
Pts: Bagley III (33)
Rebs: Bagley III (17)
Asts: Duval (11) Pts: Colson (18)
Rebs: Colson (9)
Asts: Farrell (12)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Roger Ayers, Lee Cassell, Jerry Heater
ESPN
March 8
9:00 pm
Report
No. 3 Miami 65, No. 6 North Carolina 82
Scoring by half: 31–32, 34–50
Pts: Newton (17)
Rebs: Huell, Newton (7)
Asts: Walker IV, Waardenburg (3) Pts: Pinson (25)
Rebs: Maye (13)
Asts: Johnson (5)
Barclays Center
Brooklyn, NY
Attendance: 17,732
Referees: Ron Groover, Bill Covington Jr., Raymond Styons
Semifinals[edit]
ESPN2
March 9
7:00 pm
Report
No. 1 Virginia 64, No. 4 Clemson 58
Scoring by half: 32–23, 32–35
Pts: Guy (15)
Rebs: Salt (8)
Asts: Jerome (10) Pts: Mitchell (18)
Rebs: Thomas (7)
Asts: DeVoe (3)
Barclays Center
Brooklyn, NY
Referees: Roger Aires, Bill Covington Jr., Lee Cassell
ESPN
March 9
9:00 pm
Report
No. 2 Duke 69, No. 6 North Carolina 74
Scoring by half: 31-36, 38-38
Pts: Trent, Jr. (20)
Rebs: Bagley III (13)
Asts: Duval (7) Pts: Maye (17)
Rebs: Maye (10)
Asts: Pinson (7)
Barclays Center
Brooklyn, NY
Attendance: 18,157
Referees: Bert Smith, Ted Valentine, Mike Eades
Championship[edit]
ESPN
March 10
8:30 pm
Report
No. 1 Virginia 71, No. 6 North Carolina 63
Scoring by half: 34-30, 37-33
Pts: Guy (16)
Rebs: Jerome (6)
Asts: Jerome (6) Pts: Maye (20)
Rebs: Pinson (8)
Asts: Pinson (6)
Barclays Center
Brooklyn, NY
Attendance: 18,157
Referees: Tim Nestor, Ron Groover, Jamie Luckie
Awards and honors[edit]
Tournament MVP: Kyle Guy
All-Tournament Teams:[13]
First Team
Kyle Guy, Virginia
Theo Pinson, North Carolina
Luke Maye, North Carolina
Devon Hall, Virginia
Marvin Bagley, Duke
Second Team
Ty Jerome, Virginia
Joel Berry, North Carolina
Bonzie Colson, Notre Dame
Shelton Mitchell, Clemson
Ky Bowman, Boston College
See also[edit]
ACC portal
College basketball portal
2018 ACC Women's Basketball Tournament
References[edit]
Jump up ^ Virginia wins third ACC Tournament title: ‘Hoos defeats UNC, 71-63, accessed March 11, 2018
Jump up ^ ACC loves NYC: Expect tournament to be back in Brooklyn, accessed March 11, 2018
Jump up ^ "Bracket Set For 2018 New York Life ACC Tournament". theacc.com. The Atlantic Coast Conference. March 3, 2018. Retrieved March 5, 2018.
Jump up ^ "Official Basketball Box Score Game Totals Final Statistics Syracuse vs Wake Forest" (PDF). theacc.com. The Atlantic Coast Conference. March 6, 2018. Retrieved March 7, 2018.
Jump up ^ "Official Basketball Box Score Game Totals Final Statistics Syracuse vs North Carolina" (PDF). theacc.com. The Atlantic Coast Conference. March 7, 2018. Retrieved March 8, 2018.
Jump up ^ "Official Basketball Box Score Game Totals Final Statistics Boston College vs Clemson" (PDF). theacc.com. The Atlantic Coast Conference. March 8, 2018. Retrieved March 9, 2018.
Jump up ^ "Official Basketball Box Score Game Totals Final Statistics North Carolina vs Duke" (PDF). theacc.com. The Atlantic Coast Conference. March 9, 2018. Retrieved March 10, 2018.
Jump up ^ "Official Basketball Box Score Game Totals Final Statistics North Carolina vs Virginia" (PDF). theacc.com. The Atlantic Coast Conference. March 10, 2018. Retrieved March 11, 2018.
Jump up ^ "2018 New York Life ACC Tournament" (PDF). theacc.com. The Atlantic Coast Conference. Retrieved March 5, 2018.
Jump up ^ "Eagles Soar in ACC Tournament Opener". TheACC.com. Retrieved 9 March 2018.
Jump up ^ "Irish Sweat, but Push Past Pitt". TheACC.com. Retrieved 9 March 2018.
Jump up ^ "Orange Down Deacs in First Round". TheACC.com. Retrieved 9 March 2018.
Jump up ^ "Cavaliers' Ride Continues With ACC Tourney Title". theacc.com. The Atlantic Coast Conference. March 10, 2018. Retrieved March 11, 2018.
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Implantation (human embryo)
From Wikipedia, the free encyclopedia
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This article relies largely or entirely on a single source. Relevant discussion may be found on the talk page. Please help improve this article by introducing citations to additional sources. (September 2008)
Fertilization in humans. The sperm and ovum unite through fertilization, creating a conceptus that (over the course of 8-9 days) will implant in the posterior wall of body of uterus, where it will reside over the course of 9 months.
In humans, implantation is the stage of pregnancy at which the already fertilized egg adheres to the wall of the uterus. At this stage of prenatal development, the conceptus is a blastocyst. It is by this adhesion that the fetus receives oxygen and nutrients from the mother to be able to grow.
In humans, implantation of a fertilized ovum is most likely to occur about 9 days after ovulation, ranging between 6 and 12 days.[1]
Contents
[hide]
1 Implantation window
2 Adaptation of uterus
2.1 Predecidualization
2.2 Decidualization
2.2.1 Trigger
2.2.2 Parts of decidua
2.3 Decidua throughout pregnancy
2.4 Pinopodes
2.4.1 Function in implantation
2.4.2 Function during implantation
3 Adaptation of secretions
3.1 Nourishment
3.2 Growth and implantation
4 Mechanism
4.1 Zona hatching
4.1.1 Factors
4.2 Apposition
4.2.1 Location
4.2.2 Molecular Mechanism
4.3 Adhesion
4.3.1 Communication
4.3.1.1 Proteoglycan Receptors
4.4 Invasion
4.4.1 Syncytiotrophoblasts
4.4.2 Secretions
4.4.2.1 Autocrine
4.4.2.2 Dislodging
4.4.2.3 Immunosuppressive
4.4.2.4 Decidualization
4.4.2.5 Prevention of menstruation
4.4.2.6 Other factors
5 Failure
6 See also
7 References
8 Further reading
Implantation window[edit]
The reception-ready phase of the endometrium of the uterus is usually termed the "implantation window" and lasts about 4 days. The implantation window follows around 6 days after the peak in luteinizing hormone levels. With some disparity between sources, it has been stated to occur from 7 days after ovulation until 9 days after ovulation,[2] or days 6-10 postovulation.[3] On average, it occurs during the 20th to the 23rd day after the last menstrual period.[4]
The implantation window is characterized by changes to the endometrium cells, which aid in the absorption of the uterine fluid. These changes are collectively known as the plasma membrane transformation and bring the blastocyst nearer to the endometrium and immobilize it. During this stage the blastocyst can still be eliminated by being flushed out of the uterus. Scientists have hypothesized that the hormones cause a swelling that fills the flattened out uterine cavity just prior to this stage, which may also help press the blastocyst against the endometrium.[5] The implantation window may also be initiated by other preparations in the endometrium of the uterus, both structurally and in the composition of its secretions.
Adaptation of uterus[edit]
To enable implantation, the uterus goes through changes in order to be able to receive the conceptus.
Predecidualization[edit]
The endometrium increases thickness, becomes vascularized and its glands grow to be tortuous and boosted in their secretions. These changes reach their maximum about 7 days after ovulation.
Furthermore, the surface of the endometrium produces a kind of rounded cells, which cover the whole area toward the uterine cavity. This happens about 9 to 10 days after ovulation.[6] These cells are called decidual cells, which emphasises that the whole layer of them is shed off in every menstruation if no pregnancy occurs, just as leaves of deciduous trees. The uterine glands, on the other hand, decrease in activity and degenerate already 8 to 9 days[6] after ovulation in absence of pregnancy.
The decidual cells originate from the stromal cells that are always present in the endometrium. However, the decidual cells make up a new layer, the decidua. The rest of the endometrium, in addition, expresses differences between the luminal and the basal sides. The luminal cells form the zona compacta of the endometrium, in contrast to the basalolateral zona spongiosa, which consists of the rather spongy stromal cells.[6]
Decidualization[edit]
Decidualization succeeds predecidualization if pregnancy occurs. This is an expansion of it, further developing the uterine glands, the zona compacta and the epithelium of decidual cells lining it. The decidual cells become filled with lipids and glycogen and take the polyhedral shape characteristic for decidual cells.
Trigger[edit]
It is likely that the blastocyst itself makes the main contribution to this additional growing and sustaining of the decidua. An indication of this is that decidualization occurs at a higher degree in conception cycles than in nonconception cycles.[6] Furthermore, similar changes are observed when giving stimuli mimicking the natural invasion of the embryo.[6]
Parts of decidua[edit]
The decidua can be organized into separate sections, although they have the same composition.
Decidua basalis - This is the part of the decidua which is located basalolateral to the embryo after implantation.
Decidua capsularis - Decidua capsularis grows over the embryo on the luminal side, enclosing it into the endometrium. It surrounds the embryo together with decidua basalis.
Decidua parietalis - All other decidua on the uterine surface belongs to decidua parietalis.
Decidua throughout pregnancy[edit]
After implantation the decidua remains, at least through the first trimester.[6] However, its most prominent time is during the early stages of pregnancy, during implantation. Its function as a surrounding tissue is replaced by the definitive placenta. However, some elements of the decidualization remain throughout pregnancy.[6]
The compacta and spongiosa layers are still observable beneath the decidua in pregnancy. The glands of the spongiosa layer continue to secrete during the first trimester, when they degenerate. However, before that disappearance, some glands secrete unequally much. This phenomenon of hypersecretion is called the Arias-Stella phenomenon,[6] after the pathologist Javier Arias-Stella.
Pinopodes[edit]
Main article: Pinopod
Pinopodes are small, finger-like protrusions from the endometrium. They appear between day 19 and day 21[6] of gestational age. This corresponds to a fertilization age of approximately 5 to 7 days, which corresponds well with the time of implantation. They only persist for 2 to 3 days.[6] The development of them is enhanced by progesterone but inhibited by estrogens.
Function in implantation[edit]
Pinopodes endocytose uterine fluid and macromolecules in it. By doing so, the volume of the uterus decreases, taking the walls closer to the embryoblast floating in it. Thus, the period of active pinocytes might also limit the implantation window.[6]
Function during implantation[edit]
Pinopodes continue to absorb fluid, and removes most of it during the early stages of implantation.
Adaptation of secretions[edit]
proteins, glycoproteins and peptides
secreted by the endometrial glands[6]
Matrix-associated:
Fibronectin
Laminin
Entactin
Type-IV collagen
heparan sulfate
Proteoglycan
Integrins
-
Others:
Mucins
Prolactin
IGFBP-1
Placental protein 14 (PP14) or glycodelin
Pregnancy-associated endometrial
alpha-2-globulin (alpha-2-PEG)
endometrial protein 15
Albumin
Beta-Lipoprotein
Relaxin
Fibroblast growth factor 1
Fibroblast growth factor 2
Pregnancy-associated plasma protein A
(PAPP-A)
Stress response protein 27 (SRP-27)
CA-125
Beta-endorphin
Leu-enkephalin
Diamine oxidase
Tissue plasminogen activator
Renin
Progesterone-dependent carbonic anhydrase
Lactoferrin
Not only the lining of the uterus transforms, but in addition, the secretion from its epithelial glands changes. This change is induced by increased levels of progesterone from the corpus luteum. The target of the secretions is the embryoblast, and has several functions on it.
Nourishment[edit]
The embryoblast spends approximately 72 hours[6] in the uterine cavity before implanting. In that time, it cannot receive nourishment directly from the blood of the mother, and must rely on secreted nutrients into the uterine cavity, e.g. iron[6] and fat-soluble vitamins.[6]
Growth and implantation[edit]
In addition to nourishment, the endometrium secretes several steroid-dependent proteins,[6] important for growth and implantation. Cholesterol[6] and steroids[6] are also secreted. Implantation is further facilitated by synthesis of matrix substances, adhesion molecules and surface receptors for the matrix substances. When the sperm nucleus and the egg nucleus have fused together, they form a zygote
Mechanism[edit]
Implantation is initiated when the blastocyst comes into contact with the uterine wall.
Zona hatching[edit]
Main article: Zona hatching
To be able to perform implantation, the blastocyst first needs to get rid of its zona pellucida. This process can be called "hatching".
Factors[edit]
Lytic factors in the uterine cavity, as well as factors from the blastocyst itself are essential for this process. Mechanisms in the latter are indicated by that the zona pellucida remains intact if an unfertilized egg is placed in the uterus under the same conditions.[6] A substance probably involved is plasmin. Plasminogen, the plasmin precursor, is found in the uterine cavity, and blastocyst factors contribute to its conversion to active plasmin. This hypothesis is supported by lytic effects in vitro by plasmin.[6] Furthermore, plasmin inhibitors also inhibit the entire zona hatching in rat experiments.[6]
Apposition[edit]
The very first, albeit loose, connection between the blastocyst and the endometrium is called the apposition.[6]
Location[edit]
On the endometrium, the apposition is usually made where there is a small crypt in it, perhaps because it increases the area of contact with the rather spherical blastocyst.
On the blastocyst, on the other hand, it occurs at a location where there has been enough lysis of the zona pellucida to have created a rupture to enable direct contact between the underlying trophoblast and the decidua of the endometrium.[6] However, ultimately, the inner cell mass, inside the trophoblast layer, is aligned closest to the decidua. Nevertheless, the apposition on the blastocyst is not dependent on if it is on the same side of the blastocyst as the inner cell mass. Rather, the inner cell mass rotates inside the trophoblast to align to the apposition.[6] In short, the entire surface of the blastocyst has a potential to form the apposition to the decidua.
Molecular Mechanism[edit]
The identity of the molecules on the trophoblast and the endometrial epithelia that mediate the initial interaction between the two remain unidentified. However, a number of research groups have proposed that MUC1, a member of the Mucin family of glycosylated proteins, is involved.[7] MUC1 is a transmembrane glycoprotein expressed at the apical surface of endometrial epithelial cells during the window of implantation in humans and has been shown to be differentially expressed between fertile and infertile subjects during this time.[7] MUC1 displays carbohydrate moieties on its extracellular domain that are ligands of L-selectin, a protein expressed on the surface of trophoblast cells.[8] An in vitro model of implantation developed by Genbacev et al., gave evidence to support the hypothesis that L-selectin mediates apposition of the blastocyst to the uterine epithelium by interacting with its ligands.[9]
Adhesion[edit]
Adhesion is a much stronger attachment to the endometrium than the loose apposition.
The trophoblasts adhere by penetrating the endometrium, with protrusions of trophoblast cells.
Communication[edit]
There is massive communication between the blastocyst and the endometrium at this stage. The blastocyst signals to the endometrium to adapt further to its presence, e.g. by changes in the cytoskeleton of decidual cells. This, in turn, dislodges the decidual cells from their connection to the underlying basal lamina, which enables the blastocyst to perform the succeeding invasion.[6]
This communication is conveyed by receptor-ligand-interactions, both integrin-matrix and proteoglycan ones.
Proteoglycan Receptors[edit]
Another ligand-receptor system involved in adhesion is proteoglycan receptors, found on the surface of the decidua of the uterus. Their counterparts, the proteoglycans, are found around the trophoblast cells of the blastocyst. This ligand-receptor system also is present just at the implantation window.[6]
Invasion[edit]
Invasion is an even further establishment of the blastocyst in the endometrium.
Syncytiotrophoblasts[edit]
The protrusions of trophoblast cells that adhere into the endometrium continue to proliferate and penetrate into the endometrium. As these trophoblast cells penetrate, they differentiate to become a new type of cells, syncytiotrophoblast. The prefix syn- refers to the transformation that occurs as the boundaries between these cells disappear to form a single mass of many cell nuclei (a syncytium). The rest of the trophoblasts, surrounding the inner cell mass, are hereafter called cytotrophoblasts.
Invasion continues with the syncytiotrophoblasts reaching the basal membrane beneath the decidual cells, penetrating it and further invading into the uterine stroma. Finally, the whole embryo is embedded in the endometrium. Eventually, the syncytiotrophoblasts come into contact with maternal blood and form chorionic villi. This is the initiation of forming the placenta.
Secretions[edit]
The blastocyst secretes factors for a multitude of purposes during invasion. It secretes several autocrine factors, targeting itself and stimulating it to further invade the endometrium.[6] Furthermore, secretions loosen decidual cells from each other, prevent the embryo from being rejected by the mother, trigger the final decidualization and prevent menstruation.
Autocrine[edit]
Human chorionic gonadotropin is an autocrine growth factor for the blastocyst.[6] Insulin-like growth factor 2,[6] on the other hand, stimulates the invasiveness of it.
Dislodging[edit]
The syncytiotrophoblasts dislodges decidual cells in their way, both by degradation of cell adhesion molecules linking the decidual cells together as well as degradation of the extracellular matrix between them.
Cell adhesion molecules are degraded by syncytiotrophoblast secretion of Tumor necrosis factor-alpha. This inhibits the expression of cadherins and beta-catenin.[6] Cadherins are cell adhesion molecules, and beta-catenin helps to anchor them to the cell membrane. Inhibited expression of these molecules thus loosens the connection between decidual cells, permitting the syncytotrophoblasts and the whole embryo with them to invade into the endometrium.
The extracellular matrix is degraded by serine endopeptidases and metalloproteinases. Examples of such metalloproteinases are collagenases, gelatinases and stromelysins.[6] These collagenases digest Type-I collagen, Type-II collagen, Type-III collagen, Type-VII collagen and Type-X collagen.[6] The gelatinases exist in two forms; one digesting Type-IV collagen and one digesting gelatin.[6]
Immunosuppressive[edit]
The embryo differs from the cells of the mother, and would be rejected as a parasite by the immune system of the mother if it didn't secrete immunosuppressive agents. Such agents are Platelet-activating factor, human chorionic gonadotropin, early pregnancy factor, immunosuppressive factor, Prostaglandin E2, Interleukin 1-alpha, Interleukin 6, interferon-alpha, leukemia inhibitory factor and Colony-Stimulating Factor.
Decidualization[edit]
Factors from the blastocyst also trigger the final formation of decidual cells into their proper form. In contrast, some decidual cells in the proximity of the blastocyst degenerate, providing nutrients for it.[6]
Prevention of menstruation[edit]
Human chorionic gonadotropin (hCG) not only acts as an immunosuppressive,[6] but also "notifies" the mother's body that she is pregnant, preventing menstruation by sustaining the function of the corpus luteum.
Other factors[edit]
Other factors secreted by the blastocyst are;
Embryo-derived histamine-releasing factor
Tissue plasminogen activator as well as its inhibitors
Estradiol
β1-integrins
Fibroblast growth factor
CYTL1[10]
Transforming growth factor alpha
inhibin
Failure[edit]
Implantation failure is considered to be caused by inadequate uterine receptivity in two-thirds of cases, and by problems with the embryo itself in the other third.[11]
Inadequate uterine receptivity may be caused by abnormal cytokine and hormonal signaling as well as epigenetic alterations.[12] Recurrent implantation failure is a cause of female infertility. Therefore, pregnancy rates can be improved by optimizing endometrial receptivity for implantation.[12] Evaluation of implantation markers may help to predict pregnancy outcome and detect occult implantation deficiency.[12]
Luteal support is the administration of medication, generally progestins, for the purpose of increasing the success rate of implantation and early embryogenesis, thereby complementing the function of the corpus luteum.
In women with more than 3 implantation failures in assisted reproduction, a review of several small randomized controlled studies estimated that the use of adjunct low molecular weight heparin (LMWH) improves live birth rate by approximately 80%.[13]
See also[edit]
Embryonic diapause
Blastocyst
References[edit]
Jump up ^ Wilcox AJ, Baird DD, Weinberg CR (1999). "Time of implantation of the Conceptus and loss of pregnancy". New England Journal of Medicine. 340 (23): 1796–1799. doi:10.1056/NEJM199906103402304. PMID 10362823.
Jump up ^ Xiao, Y.; Sun, X.; Yang, X.; Zhang, J.; Xue, Q.; Cai, B.; Zhou, Y. (2010). "Leukemia inhibitory factor is dysregulated in the endometrium and uterine flushing fluid of patients with adenomyosis during implantation window". Fertility and Sterility. 94 (1): 85–89. doi:10.1016/j.fertnstert.2009.03.012. PMID 19361790.
Jump up ^ Aboubakr M. Elnashar, Gamal I. Aboul-Enein. Endometrial receptivity. Middle East Fertility Society Journal, Vol. 9, No. 1, 2004, pp. 10-24
Jump up ^ 6.2 Implantation stages from embryology.ch at by the universities of Fribourg, Lausanne and Bern (Switzerland). Retrieved May, 2012
Jump up ^ "Implantation stages". Human Embryology. Online course in embryology for medicine students developed by the universities of Fribourg, Lausanne and Bern (Switzerland) with the support of the Swiss Virtual Campus. Retrieved 6 December 2011.
^ Jump up to: a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai Boron, Walter; Emile Boulpaep (2004). Medical Physiology: A Cellular And Molecular Approaoch. Oxford: Elsevier. ISBN 1-4160-2328-3. OCLC 61527528. [page needed]
^ Jump up to: a b Margarit, L.; Taylor, A.; Roberts, M. H.; Hopkins, L.; Davies, C.; Brenton, A. G.; Conlan, R. S.; Bunkheila, A.; Joels, L.; White, J. O.; Gonzalez, D. (2010). "MUC1 as a Discriminator between Endometrium from Fertile and Infertile Patients with PCOS and Endometriosis". The Journal of Clinical Endocrinology & Metabolism. 95 (12): 5320–5329. doi:10.1210/jc.2010-0603. ISSN 0021-972X.
Jump up ^ Carson, D. D.; et al., (2006). "MUC1 is a scaffold for selectin ligands in the human uterus". Front. Biosci. 11 (1): 2903. doi:10.2741/2018.
Jump up ^ Genbacev, O. D.; et al., (2003). "Trophoblast L-selectin-mediated adhesion at the maternal-fetal interface". Science. 299: 405–8. doi:10.1126/science.1079546. PMID 12532021.
Jump up ^ Ai Z., Jing W., Fang L. (2015). "Cytokine-Like Protein 1 (Cytl1) A Potential Molecular Mediator in Embryo Implantation". PLOS ONE. 11 (1): e0147424. doi:10.1371/journal.pone.0147424. PMC 4723121  . PMID 26800213. CS1 maint: Multiple names: authors list (link)
Jump up ^ Melford, S. E.; Taylor, A. H.; Konje, J. C. (2013). "Of mice and (wo)men: factors influencing successful implantation including endocannabinoids". Human Reproduction Update. 20 (3): 415–428. doi:10.1093/humupd/dmt060. ISSN 1355-4786. PMID 24306146.
^ Jump up to: a b c Cakmak, H.; Taylor, H. S. (2010). "Implantation failure: Molecular mechanisms and clinical treatment". Human Reproduction Update. 17 (2): 242–253. doi:10.1093/humupd/dmq037. PMC 3039220  . PMID 20729534.
Jump up ^ Potdar, N.; Gelbaya, T. A.; Konje, J. C.; Nardo, L. G. (2013). "Adjunct low-molecular-weight heparin to improve live birth rate after recurrent implantation failure: A systematic review and meta-analysis". Human Reproduction Update. 19 (6): 674–684. doi:10.1093/humupd/dmt032. PMID 23912476.
Further reading[edit]
"Implantation of the blastocyst..."
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