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Acrosome reaction
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Acrosome reaction on a Sea Urchin cell
During fertilization, a sperm must first fuse with the plasma membrane and then penetrate the female egg in order to fertilize it. Fusing to the egg usually causes little problem, whereas penetrating through the egg's hard shell or extracellular matrix can present more of a problem to the sperm. Therefore, sperm cells go through a process known as the acrosome reaction which is the reaction that occurs in the acrosome of the sperm as it approaches the egg. The acrosome is a cap-like structure over the anterior half of the sperm's head.
As the sperm approaches the zona pellucida of the egg, which is necessary for initiating the acrosome reaction, the membrane surrounding the acrosome fuses with the plasma membrane of the sperm's head, exposing the contents of the acrosome. The contents include surface antigens necessary for binding to the egg's cell membrane, and numerous enzymes which are responsible for breaking through the egg's tough coating and allowing fertilization to occur.[1]
Contents
[hide]
1 Variations among species
1.1 Echinoderms
1.2 Mammals
2 The process
3 In in vitro fertilization
3.1 Assessment
4 See also
5 References
6 External links
Variations among species[edit]
There are considerable species variations in the morphology and consequences of the acrosome reaction. In several species the trigger for the acrosome reaction has been identified in a layer that surrounds the egg.
Echinoderms[edit]
In some lower animal species a protuberance (the acrosomal process) forms at the apex of the sperm head, supported by a core of actin microfilaments. The membrane at the tip of the acrosomal process fuses with the egg's plasma membrane.
In some echinoderms, including starfish and sea urchins, a major portion of the exposed acrosomal content contains a protein that temporarily holds the sperm on the egg's surface.
Mammals[edit]
In mammals the acrosome reaction releases hyaluronidase and acrosin; their role in fertilization is not yet clear. The acrosomal reaction does not begin until the sperm comes into contact with the oocyte's zona pellucida. Upon coming into contact with the zona pellucida, the acrosomal enzymes begin to dissolve and the actin filament comes into contact with the zona pellucida. Once the two meet, a calcium influx occurs, causing a signaling cascade(need references). The cortical granules inside the oocyte then fuse to the outer membrane and a transient fast block reaction occurs.
It also alters a patch of pre-existing sperm plasma membrane so that it can fuse with the egg plasma membrane(need references).
A sperm penetration assay includes an acrosome reaction test that assesses how well a sperm is able to perform during the fertilization process. Sperm that are unable to properly go through the acrosome reaction will not be able to fertilize an egg. However, this problem only occurs in about 5% of men that have the test done. This test is rather expensive and provides limited information on a man's fertility.[2]
In other cases, such as in the wood mouse Apodemus sylvaticus, premature acrosome reactions have been found to cause increased motility in aggregates of spermatozoa promoting fertilization.[3]
The process[edit]
The acrosomal reaction normally takes place in the ampulla of the fallopian tube (site of fertilization) when the sperm penetrates the secondary oocyte. A few events precede the actual acrosome reaction. The sperm cell acquires a "hyperactive motility pattern" by which its flagellum produces vigorous whip-like movements that propel the sperm through the cervical canal and uterine cavity, until it reaches the isthmus of the fallopian tube. The sperm approaches the ovum in the ampulla of the fallopian tube with the help of various mechanisms, including chemotaxis. Glycoproteins on the outer surface of the sperm then bind with glycoproteins on the zona pellucida of the ovum.
Sperm that did not initiate the acrosome reaction prior to reaching to the zona pellucida are unable to penetrate the zona pellucida. Since the acrosome reaction has already occurred, sperm are then able to penetrate the zona pellucida due to mechanical action of the tail, not because of the acrosome reaction itself.[4]
The first stage is the penetration of corona radiata, by releasing hyaluronidase from the acrosome to digest cumulus cells surrounding the oocyte and exposing acrosin attached to the inner membrane of the sperm. The cumulus cells are embedded in a gel-like substance made primarily of hyaluronic acid, and developed in the ovary with the egg and support it as it grows. It is necessary for the acrosome reaction to occur before the sperm cell reaches the zona pellucida.[4]
Acrosin digests the zona pellucida and membrane of the oocyte. Part of the sperm's cell membrane then fuses with the egg cell's membrane, and the contents of the head sink into the egg. In the mouse it has been demonstrated that ZP3, one of the proteins that make up the zona pellucida, binds to a partner molecule (to the β1,4-galactosyl transferase receptors) on the sperm. This lock-and-key type mechanism is species-specific and prevents the sperm and egg of different species from fusing. The zona pellucida also releases Ca granules to prevent additional sperm from binding. There is some evidence that this binding is what triggers the acrosome to release the enzymes that allow the sperm to fuse with the egg. It is likely that a similar mechanism occurs in other mammals, but the diversity of zona proteins across species means that the relevant protein and receptor may differ.
As the sperm arrive at the egg, it is the egg that chooses the sperm and pulls it towards her.[5] The selected sperm actually tries to swim away from the egg but is tethered to the egg by female hormones. The membrane around the egg literally opens up and swallows the sperm.[6]
Upon penetration, if all is occurring normally, the process of egg-activation occurs and the oocyte is said to have become activated. This is thought to be induced by a specific protein phospholipase c zeta. It undergoes its secondary meiotic division, and the two haploid nuclei (paternal and maternal) fuse to form a zygote. In order to prevent polyspermy and minimise the possibility of producing a triploid zygote, several changes to the egg's cell membranes renders them impenetrable shortly after the first sperm enters the egg.
The aforementioned process describes the physiologically relevant events. One should however bear in mind that a certain percentage of sperm cells will undergo a spontaneous acrosome reaction without the presence of the ovum. Those cells are not able to fertilise the egg, even if they do reach it later. Other cells will spontaneously shed their acrosome during the process of apoptosis/necrosis.
In in vitro fertilization[edit]
When using intracytoplasmic sperm injection (ICSI) for IVF, the implantation rate is higher in oocytes injected with spermatozoa that have undergone acrosome reaction (~40%) vs. those injected with nonreacted spermatozoa (~10%). The implantation rate is ~25% in when injected with both reacted and nonreacted spermatozoa. The delivery rate per cycle follows the same trend.[7]
The acrosome reaction can be stimulated in vitro by substances a sperm cell may encounter naturally such as progesterone or follicular fluid, as well as the more commonly used calcium ionophore A23187.
Assessment[edit]
Birefringence microscopy,[7] flow cytometry[8] or fluorescence microscopy can be used for assessing the shedding of the acrosome or "acrosome reaction" of a sperm sample. Flow cytometry and fluorescence microscopy are usually done after staining with a fluoresceinated lectin such as FITC-PNA, FITC-PSA, FITC-ConA, or fluoresceinated antibody such as FITC-CD46.[9] The antibodies/lectins have a high specificity for different parts of the acrosomal region, and will only bind to a specific site (acrosomal content/ inner/outer membrane). If bound to a fluorescent molecule, regions where these probes have bound can be visualised. Sperm cells with artificially induced acrosome reactions may serve as positive controls.
For fluorescence microscopy a smear of washed sperm cells are made, airdried, permealized and then stained. Such a slide is then viewed under light of a wavelength that will cause the probe to fluoresce if it is bound to the acrosomal region. At least 200 cells are viewed in an arbitrary fashion and classified as either acrosome intact (fluorescing bright green) or acrosome reacted (no probe present, or only on the equatorial region). It is then expressed as a percentage of the counted cells.
For assessment with flow cytometry the washed cells are incubated with the chosen probe, (possibly washed again) and then sampled in a flow cytometer. After gating the cell population according to forward- and side-scatter the resulting data can be analysed (E.g. mean fluorescences compared). With this technique a probe for viability, like propidium iodide (PI) could also be included in order to exclude dead cells from the acrosome assessment, since many sperm cells will spontaneously lose their acrosome when they die.
See also[edit]
Cortical reaction
Hamster zona-free ovum test
ZP3
References[edit]
Jump up ^ Swiss Virtual Campus. "Chapter 4, Fertilization". Human Embryology. universities of Fribourg, Lausanne and Bern. Retrieved 18 February 2017.
Jump up ^ "Your path to fertility: Acrosome Reaction". 2007. [unreliable medical source?]
Jump up ^ Moore, Harry; Dvoráková, Katerina; Jenkins, Nicholas; Breed, William (2002). "Exceptional sperm cooperation in the wood mouse". Nature. 418 (6894): 174–7. doi:10.1038/nature00832. PMID 12110888.
^ Jump up to: a b Inoue, N; Satouh, Y; Ikawa, M; Okabe, M; Yanagimachi, R (2011). "Acrosome-reacted mouse spermatozoa recovered from the perivitelline space can fertilize other eggs". Proceedings of the National Academy of Sciences. 108 (50): 20008–11. Bibcode:2011PNAS..10820008I. doi:10.1073/pnas.1116965108. PMC 3250175  . PMID 22084105.
Jump up ^ "New Theory on How The Aggressive Egg Attracts Sperm".
Jump up ^ Yeates, Sarah E; Diamond, Sian E; Einum, Sigurd; Emerson, Brent C; Holt, William V; Gage, Matthew J. G (2013). "Cryptic Choice of Conspecific Sperm Controlled by the Impact of Ovarian Fluid on Sperm Swimming Behavior". Evolution. 67 (12): 3523–36. doi:10.1111/evo.12208. PMC 3912916  . PMID 24299405. Lay summary – ScienceDaily (August 16, 2013).
^ Jump up to: a b Gianaroli, Luca; Magli, M. Cristina; Ferraretti, Anna P; Crippa, Andor; Lappi, Michela; Capitani, Serena; Baccetti, Baccio (2010). "Birefringence characteristics in sperm heads allow for the selection of reacted spermatozoa for intracytoplasmic sperm injection". Fertility and Sterility. 93 (3): 807–13. doi:10.1016/j.fertnstert.2008.10.024. PMID 19064263.
Jump up ^ Miyazaki, R; Fukuda, M; Takeuchi, H; Itoh, S; Takada, M (2009). "Flow Cytometry to Evaluate Acrosome-Reacted Sperm". Archives of Andrology. 25 (3): 243–51. doi:10.3109/01485019008987613. PMID 2285347.
Jump up ^ Carver-Ward, J. A; Moran-Verbeek, I. M; Hollanders, J. M. G (1997). "Comparative flow cytometric analysis of the human sperm acrosome reaction using CD46 antibody and lectins". Journal of Assisted Reproduction and Genetics. 14 (2): 111–9. doi:10.1007/BF02765780. PMC 3454831  . PMID 9048242.
External links[edit]
Acrosome reaction at the US National Library of Medicine Medical Subject Headings (MeSH)
Physiology: 5/5ch8/s5ch8_21 - Essentials of Human Physiology
Animation at stanford.edu
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text:
State Emblem of India
From Wikipedia, the free encyclopedia
Jump to: navigation, search
State Emblem of India
Details
Armiger
Republic of India
Adopted
26 January 1950
Motto
Satyameva Jayate
("Truth Alone Triumphs")
The State Emblem of India, as the national emblem of India is called, is an adaptation of the Lion Capital of Ashoka at Sarnath, preserved in the Varanasi Sarnath Museum in India. A representation of Lion Capital of Ashoka was initially adopted as the emblem of the Dominion of India in December 1947.[1] The current version of the emblem was officially adopted on 26 January 1950, the day that India became a republic.[2]
Contents
[hide]
1 Usage and description
2 Emblems of national bodies
3 Historic seals and emblems
3.1 Pre-colonial India
3.2 British rule in India
3.3 Portuguese rule in India
3.4 Dominion of India
4 See also
5 References
Usage and description[edit]
The emblem forms a part of the official letterhead of the Government of India and appears on all Indian currency as well. It also functions as the national emblem of India in many places and appears prominently on Indian passports. The Ashoka Chakra (wheel) on its base features in the centre of the national flag of India.
The usage of the emblem is regulated and restricted under State Emblem of India (Prohibition of Improper Use) Act, 2005. No individual or private organisation is permitted to use the emblem for official correspondence.
The actual Sarnath capital features four Asiatic lions standing back to back, symbolizing power, courage, confidence and pride, mounted on a circular base. At the bottom is a horse and a bull, and at its centre is a wheel (Dharma chakra). The abacus is girded with a frieze of sculptures in high relief of The Lion of the North, The Horse of the West, The Bull of the South and The Elephant of the East, separated by intervening wheels, over a lotus in full bloom, exemplifying the fountainhead of life and creative inspiration. Carved from a single block of sandstone, the polished capital is crowned by the Wheel of the Law (Dharma Chakra).
In the emblem adopted by Madhav Sawhney in 1950, only three lions are visible, the fourth being hidden from view. The wheel appears in relief in the centre of the abacus, with a bull on the right and a galloping horse on the left, and outlines of Dharma Chakras on the extreme right and left. The bell-shaped lotus beneath the abacus has been omitted.[3]
Forming an integral part of the emblem is the motto inscribed below the abacus in Devanagari script: Satyameva Jayate सत्यमेव जयते (English: Truth Alone Triumphs).[4] This is a quote from Mundaka Upanishad,[5] the concluding part of the sacred Hindu Vedas.
Dinanath Bhargava was the man handpicked by legendary painter Nandalal Bose to design the National Emblem as a 21-year-old. He died at his home in Anand Nagar, Indore, on Saturday evening, the 24th December 2016. He was 89 years old.[6]
The original Lion Capital of Ashoka
Ashoka Stambha at Indian Museum, Kolkata
Early photograph.
Photograph at time of discovery.
Sarnath pillar capital of Ashoka.
Emblems of national bodies[edit]
Emblem of the Supreme Court of India
Emblem of the Central Bureau of Investigation
Emblem of the Research and Analysis Wing
Historic seals and emblems[edit]
Pre-colonial India[edit]
Seal of the Mughal Empire
Coat of arms of the British East India Company which ruled most of the subcontinent after the Marathas until 1857
British rule in India[edit]
Coat of arms of the United Kingdom, used for all official documents, publications and correspondence during the British Raj (1858-1947)
Unofficial emblem of India during the British Raj called the Star of India
Portuguese rule in India[edit]
Coat of arms of Portuguese India (1600-1935)
Lesser coat of arms of Portuguese India (1935-1961)
Greater coat of arms of Portuguese India (1935-1951)
Greater coat of arms of Portuguese India (1951-1961)
Dominion of India[edit]
Coat of arms of the United Kingdom used by the Dominion of India (August–December 1947)
Emblem used by the Dominion of India (December 1947-January 1950) [1]
Emblem of the Constituent Assembly of India
See also[edit]
Symbols of Indian states and territories
References[edit]
^ Jump up to: a b "Press Communique' - State Emblem" (PDF). Press Information Bureau of India - Archive. Archived (PDF) from the original on 24 February 2018.
Jump up ^ "State Emblem". Know India. Government of India. Retrieved 1 May 2016.
Jump up ^ "The State Emblem Of India (Prohibition Of Improper Use) Act, 2005" (PDF). 20 December 2005. p. 4. Archived from the original (PDF) on 19 March 2013. Retrieved 15 April 2012.
Jump up ^ Kamal Dey v. Union of India and State of West Bengal (Calcutta High Court 2011-07-14). Text
Jump up ^ "Rajya Sabha Parliamentary Standing Committee On Home Affairs: 116th Report on The State Emblem Of India (Prohibition Of Improper Use) Bill, 2004" (PDF). Archived from the original (PDF) on 8 March 2013.
Jump up ^ "Artist who sketched national emblem dies". Times of India. Times of India. Retrieved 16 February 2017.
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Retrieved from "https://en.wikipedia.org/w/index.php?title=State_Emblem_of_India&oldid=836043859"
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