Thrombin

F2
2c93.png
Available structures
PDBOrtholog search: PDBe RCSB
Identifiers
AliasesF2, PT, RPRGL2, THPH1, coagulation factor II, thrombin
External IDsOMIM: 176930 MGI: 88380 HomoloGene: 426 GeneCards: F2
Orthologs
SpeciesHumanMouse
Entrez
Ensembl
UniProt
RefSeq (mRNA)

NM_000506
NM_001311257

NM_010168

RefSeq (protein)

NP_000497

NP_034298

Location (UCSC)Chr 11: 46.72 – 46.74 MbChr 2: 91.63 – 91.64 Mb
PubMed search[3][4]
Wikidata
View/Edit HumanView/Edit Mouse
Schematic diagram of the blood coagulation and protein C pathways. In the blood coagulation pathway, thrombin acts to convert factor XI to XIa, VIII to VIIIa V to Va, fibrinogen to fibrin. In addition, thrombin promotes platelet activation and aggregation via activation of protease-activated receptors on the cell membrane of the platelet. Thrombin also cross over into the protein C pathway by converting protein C into APC. APC in turn converts factor V into Vi, and VIIIa into VIIIi. Finally APC activates PAR-1 and EPCR.
Role of thrombin in the blood coagulation cascade

Thrombin (EC 3.4.21.5, fibrinogenase, thrombase, thrombofort, topical, thrombin-C, tropostasin, activated blood-coagulation factor II, blood-coagulation factor IIa, factor IIa, E thrombin, beta-thrombin, gamma-thrombin) is a serine protease, an enzyme that, in humans, is encoded by the F2 gene.[5][6] Prothrombin (coagulation factor II) is proteolytically cleaved to form thrombin in the clotting process. Thrombin in turn acts as a serine protease that converts soluble fibrinogen into insoluble strands of fibrin, as well as catalyzing many other coagulation-related reactions.

History

After the description of fibrinogen and fibrin, Alexander Schmidt hypothesised the existence of an enzyme that converts fibrinogen into fibrin in 1872.[7]

Prothrombin was discovered by Pekelharing in 1894.[8][9][10]

Physiology

Synthesis

Thrombin is produced by the enzymatic cleavage of two sites on prothrombin by activated Factor X (Xa). The activity of factor Xa is greatly enhanced by binding to activated Factor V (Va), termed the prothrombinase complex. Prothrombin is produced in the liver and is co-translationally modified in a vitamin K-dependent reaction that converts 10-12 glutamic acids in the N terminus of the molecule to gamma-carboxyglutamic acid (Gla).[11] In the presence of calcium, the Gla residues promote the binding of prothrombin to phospholipid bilayers. Deficiency of vitamin K or administration of the anticoagulant warfarin inhibits the production of gamma-carboxyglutamic acid residues, slowing the activation of the coagulation cascade.

In human adults, the normal blood level of antithrombin activity has been measured to be around 1.1 units/mL. Newborn levels of thrombin steadily increase after birth to reach normal adult levels, from a level of around 0.5 units/mL 1 day after birth, to a level of around 0.9 units/mL after 6 months of life.[12]

Mechanism of action

In the blood coagulation pathway, thrombin acts to convert factor XI to XIa, VIII to VIIIa, V to Va, fibrinogen to fibrin, and XIII to XIIIa.

Factor XIIIa is a transglutaminase that catalyzes the formation of covalent bonds between lysine and glutamine residues in fibrin. The covalent bonds increase the stability of the fibrin clot. Thrombin interacts with thrombomodulin.[13][14]

As part of its activity in the coagulation cascade, thrombin also promotes platelet activation and aggregation via activation of protease-activated receptors on the cell membrane of the platelet.

Negative feedback

Thrombin bound to thrombomodulin activates protein C, an inhibitor of the coagulation cascade. The activation of protein C is greatly enhanced following the binding of thrombin to thrombomodulin, an integral membrane protein expressed by endothelial cells. Activated protein C inactivates factors Va and VIIIa. Binding of activated protein C to protein S leads to a modest increase in its activity. Thrombin is also inactivated by antithrombin, a serine protease inhibitor.

Structure

Anchoring of bovine prothrombin to the membrane through its Gla domain.[15]

The molecular weight of prothrombin is approximately 72,000 Da. The catalytic domain is released from prothrombin fragment 1.2 to create the active enzyme thrombin, which has a molecular weight of 36,000 Da. Structurally, it is a member of the large PA clan of proteases.

Prothrombin is composed of four domains; an N-terminal Gla domain, two kringle domains and a C-terminal trypsin-like serine protease domain. Factor Xa with factor V as a cofactor leads to cleavage of the Gla and two Kringle domains (forming together a fragment called fragment 1.2) and leave thrombin, consisting solely of the serine protease domain.[16]

As is the case for all serine proteases, prothrombin is converted to active thrombin by proteolysis of an internal peptide bond, exposing a new N-terminal Ile-NH3. The historic model of activation of serine proteases involves insertion of this newly formed N-terminus of the heavy chain into the β-barrel promoting the correct conformation of the catalytic residues.[17] Contrary to crystal structures of active thrombin, hydrogen-deuterium exchange mass spectrometry studies indicate that this N-terminal Ile-NH3 does not become inserted into the β-barrel in the apo form of thrombin. However, binding of the active fragment of thrombomodulin appears to allosterically promote the active conformation of thrombin by inserting this N-terminal region.[18]

Gene

The thrombin (prothrombin) gene is located on the eleventh chromosome (11p11-q12).[5]

There are an estimated 30 people in the world that have been diagnosed with the congenital form of Factor II deficiency,[19] which should not be confused with the prothrombin G20210A mutation, which is also called the factor II mutation. Prothrombin G20210A is congenital.[20]

Prothrombin G20210A is not usually accompanied by other factor mutations (i.e., the most common is factor V Leiden). The gene may be inherited heterozygous (1 pair), or much more rarely, homozygous (2 pairs), and is not related to gender or blood type. Homozygous mutations increase the risk of thrombosis more than heterozygous mutations, but the relative increased risk is not well documented. Other potential risks for thrombosis, such as oral contraceptives may be additive. The previously reported relationship of inflammatory bowel disease (i.e., Crohn's disease or ulcerative colitis) and prothrombin G20210A or factor V Leiden mutation have been contradicted by research.[21]

Role in disease

Activation of prothrombin is crucial in physiological and pathological coagulation. Various rare diseases involving prothrombin have been described (e.g., hypoprothrombinemia). Anti-prothrombin antibodies in autoimmune disease may be a factor in the formation of the lupus anticoagulant (also known as antiphospholipid syndrome). Hyperprothrombinemia can be caused by the G20210A mutation.

Thrombin, a potent vasoconstrictor and mitogen, is implicated as a major factor in vasospasm following subarachnoid hemorrhage. Blood from a ruptured cerebral aneurysm clots around a cerebral artery, releasing thrombin. This can induce an acute and prolonged narrowing of the blood vessel, potentially resulting in cerebral ischemia and infarction (stroke).

Beyond its key role in the dynamic process of thrombus formation, thrombin has a pronounced pro-inflammatory character, which may influence the onset and progression of atherosclerosis. Acting via its specific cell membrane receptors (protease activated receptors: PAR-1, PAR-3 and PAR-4), which are abundantly expressed in all arterial vessel wall constituents, thrombin has the potential to exert pro-atherogenic actions such as inflammation, leukocyte recruitment into the atherosclerotic plaque, enhanced oxidative stress, migration and proliferation of vascular smooth muscle cells, apoptosis and angiogenesis.[22][23][24]

Thrombin is implicated in the physiology of blood clots. Its presence indicates the existence of a clot. In 2013 a system for detecting the presence of thrombin was developed in mice. It combines peptide-coated iron oxide attached to "reporter chemicals". When a peptide binds to a thrombin molecule, the report is released and appears in the urine where it can be detected. Human testing has not been conducted.[25]

Applications

Research tool

Due to its high proteolytic specificity, thrombin is a valuable biochemical tool. The thrombin cleavage site (Leu-Val-Pro-Arg-Gly-Ser) is commonly included in linker regions of recombinant fusion protein constructs. Following purification of the fusion protein, thrombin can be used to selectively cleave between the arginine and glycine residues of the cleavage site, effectively removing the purification tag from the protein of interest with a high degree of specificity.

Medicine and surgery

Prothrombin complex concentrate and fresh frozen plasma are prothrombin-rich coagulation factor preparations that can be used to correct deficiencies (usually due to medication) of prothrombin. Indications include intractable bleeding due to warfarin.

Manipulation of prothrombin is central to the mode of action of most anticoagulants. Warfarin and related drugs inhibit vitamin K-dependent carboxylation of several coagulation factors, including prothrombin. Heparin increases the affinity of antithrombin to thrombin (as well as factor Xa). The direct thrombin inhibitors, a newer class of medication, directly inhibit thrombin by binding to its active site.

Recombinant thrombin is available as a powder for reconstitution into aqueous solution. It can be applied topically during surgery, as an aid to hemostasis. It can be useful for controlling minor bleeding from capillaries and small venules, but ineffective and not indicated for massive or brisk arterial bleeding.[26][27][28]

Food production

Thrombin, combined with fibrinogen, is sold under the brand name Fibrimex for use as a binding agent for meat. Both proteins in Fibrimex derives from porcine or bovine blood.[29] According to the manufacturer it can be used to produce new kinds of mixed meats (for example combining beef and fish seamlessly). The manufacturer also states that it can be used to combine whole muscle meat, form and portion these, thus cutting down on production costs without a loss in quality.[30]

General secretary Jan Bertoft of Swedish Consumers' Association has stated that "there is danger of misleading the consumers since there is no way to tell this reconstituted meat from real meat".[29]

See also

References

  1. ^ a b c GRCh38: Ensembl release 89: ENSG00000180210 - Ensembl, May 2017
  2. ^ a b c GRCm38: Ensembl release 89: ENSMUSG00000027249 - Ensembl, May 2017
  3. ^ "Human PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
  4. ^ "Mouse PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
  5. ^ a b Royle NJ, Irwin DM, Koschinsky ML, MacGillivray RT, Hamerton JL (May 1987). "Human genes encoding prothrombin and ceruloplasmin map to 11p11-q12 and 3q21-24, respectively". Somatic Cell and Molecular Genetics. 13 (3): 285–92. doi:10.1007/BF01535211. PMID 3474786. S2CID 45686258.
  6. ^ Degen SJ, Davie EW (September 1987). "Nucleotide sequence of the gene for human prothrombin". Biochemistry. 26 (19): 6165–77. doi:10.1021/bi00393a033. PMID 2825773.
  7. ^ Schmidt A (1872). "Neue Untersuchungen ueber die Fasserstoffesgerinnung". Pflügers Archiv für die gesamte Physiologie. 6: 413–538. doi:10.1007/BF01612263. S2CID 37273997.
  8. ^ Kaushansky K, Lichtman M, Prchal J, Levi M, Press O, Burns L, Caligiuri M (2015). Williams Hematology. McGraw-Hill. p. 1918.
  9. ^ Quick AJ (1957). Hemorrhagic Diseases. Philadelphia: Lea and Febiger. pp. 451–490.
  10. ^ Morawitz P (1905). "Die Chemie der Blutgerinnung". Ergeb Physiol. 4: 307–422. doi:10.1007/BF02321003. S2CID 84003009.
  11. ^ Knorre DG, Kudryashova NV, Godovikova TS (October 2009). "Chemical and functional aspects of posttranslational modification of proteins". Acta Naturae. 1 (3): 29–51. doi:10.32607/20758251-2009-1-3-29-51. PMC 3347534. PMID 22649613.
  12. ^ Andrew M, Paes B, Milner R, Johnston M, Mitchell L, Tollefsen DM, Powers P (July 1987). "Development of the human coagulation system in the full-term infant". Blood. 70 (1): 165–72. doi:10.1182/blood.V70.1.165.165. PMID 3593964.
  13. ^ Bajzar L, Morser J, Nesheim M (July 1996). "TAFI, or plasma procarboxypeptidase B, couples the coagulation and fibrinolytic cascades through the thrombin-thrombomodulin complex". The Journal of Biological Chemistry. 271 (28): 16603–8. doi:10.1074/jbc.271.28.16603. PMID 8663147.
  14. ^ Jakubowski HV, Owen WG (July 1989). "Macromolecular specificity determinants on thrombin for fibrinogen and thrombomodulin". The Journal of Biological Chemistry. 264 (19): 11117–21. doi:10.1016/S0021-9258(18)60437-5. PMID 2544585.
  15. ^ PDB: 1nl2​;Huang M, Rigby AC, Morelli X, Grant MA, Huang G, Furie B, Seaton B, Furie BC (September 2003). "Structural basis of membrane binding by Gla domains of vitamin K-dependent proteins". Nature Structural Biology. 10 (9): 751–6. doi:10.1038/nsb971. PMID 12923575. S2CID 7751100.
  16. ^ Davie EW, Kulman JD (April 2006). "An overview of the structure and function of thrombin". Seminars in Thrombosis and Hemostasis. 32 Suppl 1: 3–15. doi:10.1055/s-2006-939550. PMID 16673262.
  17. ^ Huber R, Bode W (1978-03-01). "Structural basis of the activation and action of trypsin". Accounts of Chemical Research. 11 (3): 114–122. doi:10.1021/ar50123a006. ISSN 0001-4842.
  18. ^ Handley LD, Treuheit NA, Venkatesh VJ, Komives EA (November 2015). "Thrombomodulin Binding Selects the Catalytically Active Form of Thrombin". Biochemistry. 54 (43): 6650–8. doi:10.1021/acs.biochem.5b00825. PMC 4697735. PMID 26468766.
  19. ^ Degen SJ, McDowell SA, Sparks LM, Scharrer I (February 1995). "Prothrombin Frankfurt: a dysfunctional prothrombin characterized by substitution of Glu-466 by Ala". Thrombosis and Haemostasis. 73 (2): 203–9. doi:10.1055/s-0038-1653751. PMID 7792730.
  20. ^ Varga EA, Moll S (July 2004). "Cardiology patient pages. Prothrombin 20210 mutation (factor II mutation)". Circulation. 110 (3): e15–8. doi:10.1161/01.CIR.0000135582.53444.87. PMID 15262854.
  21. ^ Bernstein CN, Sargent M, Vos HL, Rosendaal FR (February 2007). "Mutations in clotting factors and inflammatory bowel disease". The American Journal of Gastroenterology. 102 (2): 338–43. PMID 17156138.
  22. ^ Borissoff JI, Spronk HM, Heeneman S, ten Cate H (June 2009). "Is thrombin a key player in the 'coagulation-atherogenesis' maze?". Cardiovascular Research. 82 (3): 392–403. doi:10.1093/cvr/cvp066. PMID 19228706.
  23. ^ Borissoff JI, Heeneman S, Kilinç E, Kassák P, Van Oerle R, Winckers K, Govers-Riemslag JW, Hamulyák K, Hackeng TM, Daemen MJ, ten Cate H, Spronk HM (August 2010). "Early atherosclerosis exhibits an enhanced procoagulant state". Circulation. 122 (8): 821–30. doi:10.1161/CIRCULATIONAHA.109.907121. PMID 20697022.
  24. ^ Borissoff JI, Spronk HM, ten Cate H (May 2011). "The hemostatic system as a modulator of atherosclerosis". The New England Journal of Medicine. 364 (18): 1746–60. doi:10.1056/NEJMra1011670. PMID 21542745.
  25. ^ Economist (2013-11-05). "Nanomedicine: Particle physiology". The Economist. Retrieved 2013-12-15.
  26. ^ Chapman WC, Singla N, Genyk Y, McNeil JW, Renkens KL, Reynolds TC, Murphy A, Weaver FA (August 2007). "A phase 3, randomized, double-blind comparative study of the efficacy and safety of topical recombinant human thrombin and bovine thrombin in surgical hemostasis". Journal of the American College of Surgeons. 205 (2): 256–65. doi:10.1016/j.jamcollsurg.2007.03.020. PMID 17660072.
  27. ^ Singla NK, Ballard JL, Moneta G, Randleman CD, Renkens KL, Alexander WA (July 2009). "A phase 3b, open-label, single-group immunogenicity and safety study of topical recombinant thrombin in surgical hemostasis". Journal of the American College of Surgeons. 209 (1): 68–74. doi:10.1016/j.jamcollsurg.2009.03.016. PMID 19651065.
  28. ^ Greenhalgh DG, Gamelli RL, Collins J, Sood R, Mozingo DW, Gray TE, Alexander WA (2009). "Recombinant thrombin: safety and immunogenicity in burn wound excision and grafting". Journal of Burn Care & Research. 30 (3): 371–9. doi:10.1097/BCR.0b013e3181a28979. PMID 19349898. S2CID 3678462.
  29. ^ a b "Sverige röstade ja till köttklister" [Sweden voted in favor of the meat paste] (in Swedish). Dagens Nyheter. 2010-02-09. Retrieved 2010-10-17.
  30. ^ "Welcome to Fibrimex". Fibrimex website. Sonac. Retrieved 2019-02-28.

Further reading

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Crystal structure of thrombin with an inhibitor. Created with pymol from PDB 2c93
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Cartoon representation of the molecular structure of protein registered with 1a61 code.
PDB 1h8d EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1h8d code.
PDB 1c1w EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1c1w code.
PDB 1a2c EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1a2c code.
PDB 1g30 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1g30 code.
PDB 1c1u EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1c1u code.
PDB 1tom EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1tom code.
PDB 1vr1 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1vr1 code.
PDB 1qj6 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1qj6 code.
Ideogram human chromosome 11.svg
Ideogram of human chromosome. Chromosome 11 highlighted. G-band, 850 bphs (bands per haploid set). Black and gray: Giemsa positive. Red: Centromere. Light blue: Variable region. Dark blue: Stalk.
PDB 1dwe EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1dwe code.
PDB 1way EBI.png
Cartoon representation of the molecular structure of protein registered with 1way code.
Ideogram of house mouse chromosome 2.svg
House mouse (Mus musculus) chromosome 2.
PDB 1nzq EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1nzq code.
PDB 1bhx EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1bhx code.
PDB 1hao EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1hao code.
PDB 1t4v EBI.png
Cartoon representation of the molecular structure of protein registered with 1t4v code.
PDB 1o5g EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1o5g code.
PDB 1xmn EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1xmn code.
PDB 1p8v EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1p8v code.
PDB 2h9t EBI.png
Cartoon representation of the molecular structure of protein registered with 2h9t code.
PDB 2c90 EBI.png
Cartoon representation of the molecular structure of protein registered with 2c90 code.
PDB 1b7x EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1b7x code.
PDB 2hgt EBI.jpg
Cartoon representation of the molecular structure of protein registered with 2hgt code.
PDB 2od3 EBI.png
Cartoon representation of the molecular structure of protein registered with 2od3 code.
PDB 1tmt EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1tmt code.
PDB 2bvs EBI.png
Cartoon representation of the molecular structure of protein registered with 2bvs code.
PDB 1nrs EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1nrs code.
PDB 1ad8 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ad8 code.
PDB 1tmb EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1tmb code.
PDB 1bbr EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1bbr code.
PDB 1c5l EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1c5l code.
PDB 1awf EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1awf code.
PDB 1sl3 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1sl3 code.
PDB 1eol EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1eol code.
PDB 1ypl EBI.png
Cartoon representation of the molecular structure of protein registered with 1ypl code.
PDB 2jh0 EBI.png
Cartoon representation of the molecular structure of protein registered with 2jh0 code.
PDB 1tbz EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1tbz code.
PDB 1vit EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1vit code.
PDB 1hgt EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1hgt code.
PDB 1a5g EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1a5g code.
PDB 1ai8 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ai8 code.
PDB 1aix EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1aix code.
PDB 1ook EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ook code.
PDB 1c4v EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1c4v code.
PDB 1hlt EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1hlt code.
PDB 1dit EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1dit code.
PDB 1d3d EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1d3d code.
PDB 1ypk EBI.png
Cartoon representation of the molecular structure of protein registered with 1ypk code.
PDB 7kme EBI.jpg
Cartoon representation of the molecular structure of protein registered with 7kme code.
PDB 1twx EBI.png
Cartoon representation of the molecular structure of protein registered with 1twx code.
PDB 2bxt EBI.png
Cartoon representation of the molecular structure of protein registered with 2bxt code.
PDB 2jh6 EBI.png
Cartoon representation of the molecular structure of protein registered with 2jh6 code.
PDB 1ghv EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ghv code.
PDB 1t4u EBI.png
Cartoon representation of the molecular structure of protein registered with 1t4u code.
PDB 1gj5 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1gj5 code.
PDB 2c8x EBI.png
Cartoon representation of the molecular structure of protein registered with 2c8x code.
PDB 1mue EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1mue code.
PDB 2cf8 EBI.png
Cartoon representation of the molecular structure of protein registered with 2cf8 code.
PDB 1ay6 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ay6 code.
PDB 1ucy EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ucy code.
PDB 1d3p EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1d3p code.
PDB 2bvx EBI.png
Cartoon representation of the molecular structure of protein registered with 2bvx code.
PDB 1sgi EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1sgi code.
PDB 2anm EBI.png
Cartoon representation of the molecular structure of protein registered with 2anm code.
PDB 1ypj EBI.png
Cartoon representation of the molecular structure of protein registered with 1ypj code.
PDB 8kme EBI.jpg
Cartoon representation of the molecular structure of protein registered with 8kme code.
PDB 1d3t EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1d3t code.
PDB 1sb1 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1sb1 code.
PDB 2b5t EBI.jpg
Cartoon representation of the molecular structure of protein registered with 2b5t code.
PDB 1nrp EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1nrp code.
PDB 1sr5 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1sr5 code.
PDB 1uvs EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1uvs code.
PDB 2c8z EBI.png
Cartoon representation of the molecular structure of protein registered with 2c8z code.
PDB 1ta2 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ta2 code.
PDB 1lhe EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1lhe code.
PDB 1ype EBI.png
Cartoon representation of the molecular structure of protein registered with 1ype code.
PDB 1b5g EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1b5g code.
PDB 1hxe EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1hxe code.
PDB 1qhr EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1qhr code.
PDB 1hap EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1hap code.
PDB 1dm4 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1dm4 code.
PDB 1ypm EBI.png
Cartoon representation of the molecular structure of protein registered with 1ypm code.
PDB 1d6w EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1d6w code.
PDB 1riw EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1riw code.
PDB 1nrq EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1nrq code.
PDB 1jou EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1jou code.
PDB 2a45 EBI.png
Cartoon representation of the molecular structure of protein registered with 2a45 code.
PDB 1a4w EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1a4w code.
PDB 1ny2 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ny2 code.
PDB 1dx5 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1dx5 code.
PDB 2cn0 EBI.png
Cartoon representation of the molecular structure of protein registered with 2cn0 code.
PDB 2cf9 EBI.png
Cartoon representation of the molecular structure of protein registered with 2cf9 code.
PDB 1ae8 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ae8 code.
PDB 2jh5 EBI.png
Cartoon representation of the molecular structure of protein registered with 2jh5 code.
PDB 1k21 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1k21 code.
PDB 1gj4 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1gj4 code.
PDB 1rd3 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1rd3 code.
Ideogram house mouse chromosome 2.svg
Ideogram of house mouse (Mus musculus) chromosome. Chromosome 2 highlighted.
PDB 2bvr EBI.png
Cartoon representation of the molecular structure of protein registered with 2bvr code.
PDB 1ta6 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ta6 code.
PDB 1abj EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1abj code.
PDB 3hat EBI.jpg
Cartoon representation of the molecular structure of protein registered with 3hat code.
PDB 1hai EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1hai code.
PDB 1c4u EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1c4u code.
PDB 1qj7 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1qj7 code.
PDB 1ca8 EBI.jpg
Cartoon representation of the molecular structure of protein registered with 1ca8 code.
Human chromosome 11 ideogram.svg
Human chromosome 11. G-banding ideogram in resolution 850 bphs.