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{{protein
{{Infobox_gene}}
|Name=uncoupling protein 1 (mitochondrial, proton carrier)
'''Thermogenin''' (called [[uncoupling protein]] by its discoverers and now known as uncoupling protein 1, or '''UCP1''')<ref name="entrez">{{cite web | title = Entrez Gene: UCP1 uncoupling protein 1 (mitochondrial, proton carrier)| url = https://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=7350| accessdate = }}</ref> is an uncoupling protein found in the [[Mitochondrion|mitochondria]] of [[brown adipose tissue]] (BAT). It is used to generate heat by non-shivering [[thermogenesis]], and makes a quantitatively important contribution to countering heat loss in [[neonates]] which would otherwise occur due to their high surface area-volume ratio.
|caption=
|image=
|width=
|HGNCid=12517
|Symbol=UCP1
|AltSymbols=UCP
|EntrezGene=7350
|OMIM=113730
|RefSeq=NM_021833
|UniProt=P25874
|PDB=
|ECnumber=
|Chromosome=4
|Arm=q
|Band=31
|LocusSupplementaryData=
}}
{{SI}}
 
 
==Overview==
'''Thermogenin''' (called [[uncoupling protein]] by its discoverers and now known as uncoupling protein 1, or '''UCP1''') is an uncoupling protein found in the [[Mitochondrion|mitochondria]] of [[brown adipose tissue]] (BAT). It is used to generate heat by non-shivering [[thermogenesis]].  Non-shivering thermogenesis is the primary means of heat generation in hibernating [[mammals]] and in human infants.


==Mechanism==
==Mechanism==
The molecular mechanism of UCP1 mediated uncoupling is reasonably well understood; UCP1 provides an alternative pathway by which protons can reenter the mitochondrial matrix, short-circuiting the 'proton circuit' linking the respiratory chain to the ATP synthase (which generates ATP for the cell) and allowing respiration (and hence heat production) to proceed in the absence of ATP synthesis. UCP1 is related to other mitochondrial metabolite transporters such as the adenine nucleotide translocator a proton channel in the [[mitochondrial inner membrane]] that permits the translocation of protons from the mitochondrial [[intermembrane space]] to the [[mitochondrial matrix]]. UCP1 is restricted to brown fat where it provides a mechanism for the enormous heat generating capacity of the tissue.
[[File:ThermogeneseAdipozyten-en.svg|thumb|left|Mechanism of thermogenin activation: In a last step thermogenin inhibition is released through the presence of free fatty acids. The cascade is initiated by binding of norepinephrine to the cells β<sub>3</sub>-adrenoceptors.]]
UCPs are transmembrane proteins that decrease the proton gradient generated in oxidative phosphorylation. They do this by increasing the permeability of the inner mitochondrial membrane, allowing protons that have been pumped into the intermembrane space to return to the mitochondrial matrix. UCP1-mediated heat generation in brown fat uncouples the respiratory chain, allowing for fast substrate oxidation with a low rate of ATP production. UCP1 is related to other mitochondrial metabolite transporters such as the adenine nucleotide translocator, a proton channel in the [[mitochondrial inner membrane]] that permits the translocation of protons from the mitochondrial [[intermembrane space]] to the [[mitochondrial matrix]]. UCP1 is restricted to [[brown adipose tissue]], where it provides a mechanism for the enormous heat-generating capacity of the tissue.


UCP1 is activated in the brown fat cell by fatty acids that are liberated by the following pathway: Sympathetic nervous system terminals release noradrenaline onto a beta3 membrane receptor. This activates [[adenylyl cyclase]] which catalyses the conversion of ATP to [[cyclic AMP]] (cAMP). cAMP activates protein kinase A causing its active C subunits to be freed from its regulatory R subunits. Active protein kinase A in turn phosphorylates [[triacylglycerol lipase]], thereby activating it. The lipase converts triacylglycerols into free fatty acids which activate UCP1, overriding the inhibition causes by cytoplasmic ATP. At the termination of thermogenesis, the mitochondria oxidize away the residual fatty acids, UCP1 inactivates and the cell resumes its normal energy-conserving mode.
UCP1 is activated in the brown fat cell by fatty acids and inhibited by nucleotides. Fatty acids cause the following signaling cascade: Sympathetic nervous system terminals release [[Norepinephrine]] onto a [[Beta-3 adrenergic receptor]] on the [[plasma membrane]]. This activates [[adenylyl cyclase]], which catalyses the conversion of ATP to [[cyclic AMP]] (cAMP). cAMP activates [[protein kinase A]], causing its active C subunits to be freed from its regulatory R subunits. Active protein kinase A, in turn, phosphorylates [[triacylglycerol lipase]], thereby activating it. The lipase converts triacylglycerols into free fatty acids, which activate UCP1, overriding the inhibition caused by purine nucleotides ([[Guanosine diphosphate|GDP]] and [[Adenosine diphosphate|ADP]]). At the termination of thermogenesis, the mitochondria oxidize away the residual fatty acids, UCP1 inactivates and the cell resumes its normal energy-conserving mode.


==History==
==History==
Uncoupling protein 1 was discovered in 1979 - see <ref>{{cite journal |author=Nicholls D, Bernson V, Heaton G |title=The identification of the component in the inner membrane of brown adipose tissue mitochondria responsible for regulating energy dissipation |journal=Experientia Suppl |volume=32 |issue= |pages=89-93 |year= |pmid=348493}}</ref> and was first cloned in 1988.<ref>http://www.jbc.org/cgi/reprint/263/25/12274</ref><ref>
Uncoupling protein 1 was discovered in 1978<ref name="pmid348493">{{cite journal | vauthors = Nicholls DG, Bernson VS, Heaton GM | title = The identification of the component in the inner membrane of brown adipose tissue mitochondria responsible for regulating energy dissipation | journal = Experientia. Supplementum | volume = 32 | issue = | pages = 89–93 | year = 1978 | pmid = 348493 | doi = 10.1007/978-3-0348-5559-4_9 }}</ref> and was first cloned in 1988.<ref name="pmid3410843">{{cite journal | vauthors = Kozak LP, Britton JH, Kozak UC, Wells JM | title = The mitochondrial uncoupling protein gene. Correlation of exon structure to transmembrane domains | journal = The Journal of Biological Chemistry | volume = 263 | issue = 25 | pages = 12274–7 | date = Sep 1988 | pmid = 3410843 | doi =  | url = http://www.jbc.org/cgi/content/abstract/263/25/12274 }}</ref><ref name="pmid3202878">{{cite journal | vauthors = Bouillaud F, Raimbault S, Ricquier D | title = The gene for rat uncoupling protein: complete sequence, structure of primary transcript and evolutionary relationship between exons | journal = Biochemical and Biophysical Research Communications | volume = 157 | issue = 2 | pages = 783–92 | date = Dec 1988 | pmid = 3202878 | doi = 10.1016/S0006-291X(88)80318-8 }}</ref>
 
[[Uncoupling protein two]] (UCP2), a homolog of UCP1, was identified in 1997. UCP2 localizes to a wide variety of tissues, and is thought to be involved in regulating reactive oxygen species (ROS). In the past decade, three additional homologs of UCP1 have been identified, including UCP3, UCP4, and BMCP1 (also known as UCP5).


{{cite journal |author=Bouillaud F, Raimbault S, Ricquier D |title=The gene for rat uncoupling protein: complete sequence, structure of primary transcript and evolutionary relationship between exons |journal=Biochem Biophys Res Commun |volume=157 |issue=2 |pages=783-92 |year=1988 |pmid=3202878}} [http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WBK-4G3D5KD-4V&_coverDate=12%2F15%2F1988&_alid=424246349&_rdoc=1&_fmt=&_orig=search&_qd=1&_cdi=6713&_sort=d&view=c&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=adaf6622e66b1b6d02808d2f81bfbcab link]</ref>  
== Clinical relevance ==
Methods of delivering UCP1 to cells by gene transfer therapy or methods of its upregulation have been an important line of enquiry in research into the treatment of obesity, due to their ability to dissipate excess metabolic stores.<ref>{{cite journal | vauthors = Kozak LP, Anunciado-Koza R | title = UCP1: its involvement and utility in obesity | journal = International Journal of Obesity | volume = 32 Suppl 7 | issue = Suppl 7 | pages = S32-8 | date = Dec 2008 | pmid = 19136989 | pmc = 2746324 | doi = 10.1038/ijo.2008.236 }}</ref>


[[Uncoupling protein two]] (UCP2), a homolog of UCP1, was identified in 1997. 
== See also ==
* [[2,4-Dinitrophenol]] (A synthetic [[small-molecule]] proton shuttle with similar effects)


In the past decade three additional homologs of UCP1 have been identified, including UCP3, UCP4, and BMCP1 (also known as UCP5).
== References ==
{{Reflist|2}}


==References==
== Further reading ==
<references/>
{{refbegin | 2}}
* {{cite journal | vauthors = Ricquier D, Bouillaud F | title = The uncoupling protein homologues: UCP1, UCP2, UCP3, StUCP and AtUCP | journal = The Biochemical Journal | volume = 345 Pt 2 | issue = 2 | pages = 161–79 | date = Jan 2000 | pmid = 10620491 | pmc = 1220743 | doi = 10.1042/0264-6021:3450161 }}
* {{cite journal | vauthors = Muzzin P | title = The uncoupling proteins | journal = Annales d'Endocrinologie | volume = 63 | issue = 2 Pt 1 | pages = 106–10 | date = Apr 2002 | pmid = 11994670 | doi =  }}
* {{cite journal | vauthors = Del Mar Gonzalez-Barroso M, Ricquier D, Cassard-Doulcier AM | title = The human uncoupling protein-1 gene (UCP1): present status and perspectives in obesity research | journal = Obesity Reviews | volume = 1 | issue = 2 | pages = 61–72 | date = Oct 2000 | pmid = 12119988 | doi =  10.1046/j.1467-789x.2000.00009.x}}
* {{cite journal | vauthors = Cassard AM, Bouillaud F, Mattei MG, Hentz E, Raimbault S, Thomas M, Ricquier D | title = Human uncoupling protein gene: structure, comparison with rat gene, and assignment to the long arm of chromosome 4 | journal = Journal of Cellular Biochemistry | volume = 43 | issue = 3 | pages = 255–64 | date = Jul 1990 | pmid = 2380264 | doi = 10.1002/jcb.240430306 }}
* {{cite journal | vauthors = Bouillaud F, Villarroya F, Hentz E, Raimbault S, Cassard AM, Ricquier D | title = Detection of brown adipose tissue uncoupling protein mRNA in adult patients by a human genomic probe | journal = Clinical Science | volume = 75 | issue = 1 | pages = 21–7 | date = Jul 1988 | pmid = 3165741 | doi =  10.1042/cs0750021}}
* {{cite journal | vauthors = Oppert JM, Vohl MC, Chagnon M, Dionne FT, Cassard-Doulcier AM, Ricquier D, Pérusse L, Bouchard C | title = DNA polymorphism in the uncoupling protein (UCP) gene and human body fat | journal = International Journal of Obesity and Related Metabolic Disorders | volume = 18 | issue = 8 | pages = 526–31 | date = Aug 1994 | pmid = 7951471 | doi =  }}
* {{cite journal | vauthors = Clément K, Ruiz J, Cassard-Doulcier AM, Bouillaud F, Ricquier D, Basdevant A, Guy-Grand B, Froguel P | title = Additive effect of A-->G (-3826) variant of the uncoupling protein gene and the Trp64Arg mutation of the beta 3-adrenergic receptor gene on weight gain in morbid obesity | journal = International Journal of Obesity and Related Metabolic Disorders | volume = 20 | issue = 12 | pages = 1062–6 | date = Dec 1996 | pmid = 8968850 | doi =  }}
* {{cite journal | vauthors = Schleiff E, Shore GC, Goping IS | title = Human mitochondrial import receptor, Tom20p. Use of glutathione to reveal specific interactions between Tom20-glutathione S-transferase and mitochondrial precursor proteins | journal = FEBS Letters | volume = 404 | issue = 2-3 | pages = 314–8 | date = Mar 1997 | pmid = 9119086 | doi = 10.1016/S0014-5793(97)00145-2 }}
* {{cite journal | vauthors = Urhammer SA, Fridberg M, Sørensen TI, Echwald SM, Andersen T, Tybjaerg-Hansen A, Clausen JO, Pedersen O | title = Studies of genetic variability of the uncoupling protein 1 gene in Caucasian subjects with juvenile-onset obesity | journal = The Journal of Clinical Endocrinology and Metabolism | volume = 82 | issue = 12 | pages = 4069–74 | date = Dec 1997 | pmid = 9398715 | doi = 10.1210/jc.82.12.4069 }}
* {{cite journal | vauthors = Jezek P, Urbánková E | title = Specific sequence of motifs of mitochondrial uncoupling proteins | journal = IUBMB Life | volume = 49 | issue = 1 | pages = 63–70 | date = Jan 2000 | pmid = 10772343 | doi = 10.1080/713803586 }}
* {{cite journal | vauthors = Mori H, Okazawa H, Iwamoto K, Maeda E, Hashiramoto M, Kasuga M | title = A polymorphism in the 5' untranslated region and a Met229-->Leu variant in exon 5 of the human UCP1 gene are associated with susceptibility to type II diabetes mellitus | journal = Diabetologia | volume = 44 | issue = 3 | pages = 373–6 | date = Mar 2001 | pmid = 11317671 | doi = 10.1007/s001250051629 }}
* {{cite journal | vauthors = Nibbelink M, Moulin K, Arnaud E, Duval C, Pénicaud L, Casteilla L | title = Brown fat UCP1 is specifically expressed in uterine longitudinal smooth muscle cells | journal = The Journal of Biological Chemistry | volume = 276 | issue = 50 | pages = 47291–5 | date = Dec 2001 | pmid = 11572862 | doi = 10.1074/jbc.M105658200 }}
* {{cite journal | vauthors = Echtay KS, Roussel D, St-Pierre J, Jekabsons MB, Cadenas S, Stuart JA, Harper JA, Roebuck SJ, Morrison A, Pickering S, Clapham JC, Brand MD | title = Superoxide activates mitochondrial uncoupling proteins | journal = Nature | volume = 415 | issue = 6867 | pages = 96–9 | date = Jan 2002 | pmid = 11780125 | doi = 10.1038/415096a }}
* {{cite journal | vauthors = Rousset S, del Mar Gonzalez-Barroso M, Gelly C, Pecqueur C, Bouillaud F, Ricquier D, Cassard-Doulcier AM | title = A new polymorphic site located in the human UCP1 gene controls the in vitro binding of CREB-like factor | journal = International Journal of Obesity and Related Metabolic Disorders | volume = 26 | issue = 5 | pages = 735–8 | date = May 2002 | pmid = 12032762 | doi = 10.1038/sj.ijo.0801973 }}
* {{cite journal | vauthors = Rim JS, Kozak LP | title = Regulatory motifs for CREB-binding protein and Nfe2l2 transcription factors in the upstream enhancer of the mitochondrial uncoupling protein 1 gene | journal = The Journal of Biological Chemistry | volume = 277 | issue = 37 | pages = 34589–600 | date = Sep 2002 | pmid = 12084707 | doi = 10.1074/jbc.M108866200 }}
* {{cite journal | vauthors = Kieć-Wilk B, Wybrańska I, Malczewska-Malec M, Leszczyńska-Gołabek L, Partyka L, Niedbał S, Jabrocka A, Dembińska-Kieć A | title = Correlation of the -3826A >G polymorphism in the promoter of the uncoupling protein 1 gene with obesity and metabolic disorders in obese families from southern Poland | journal = Journal of Physiology and Pharmacology | volume = 53 | issue = 3 | pages = 477–90 | date = Sep 2002 | pmid = 12375583 | doi =  }}
{{refend}}


==External links==
== External links ==
* [http://news.bbc.co.uk/2/hi/health/5335176.stm Seaweed anti-obesity tablet hope] (BBC - Thermogenin mentioned as part of process)
* [http://news.bbc.co.uk/2/hi/health/5335176.stm Seaweed anti-obesity tablet hope] (BBC - Thermogenin mentioned as part of process)
* {{MeshName|thermogenin}}
* {{MeshName|thermogenin}}


[[Category:Cellular respiration]]
[[Category:Cellular respiration]]
[[Category:Uncoupling Proteins]]
[[Category:Mitochondria]]
 
[[ca:Termogenina]]
[[de:Thermogenin]]
[[pl:Termogenina]]
 
{{WH}}
{{WS}}

Revision as of 17:28, 22 September 2017

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Identifiers
Aliases
External IDsGeneCards: [1]
Orthologs
SpeciesHumanMouse
Entrez

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Ensembl

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UniProt

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RefSeq (mRNA)

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RefSeq (protein)

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Location (UCSC)n/an/a
PubMed searchn/an/a
Wikidata
View/Edit Human

Thermogenin (called uncoupling protein by its discoverers and now known as uncoupling protein 1, or UCP1)[1] is an uncoupling protein found in the mitochondria of brown adipose tissue (BAT). It is used to generate heat by non-shivering thermogenesis, and makes a quantitatively important contribution to countering heat loss in neonates which would otherwise occur due to their high surface area-volume ratio.

Mechanism

File:ThermogeneseAdipozyten-en.svg
Mechanism of thermogenin activation: In a last step thermogenin inhibition is released through the presence of free fatty acids. The cascade is initiated by binding of norepinephrine to the cells β3-adrenoceptors.

UCPs are transmembrane proteins that decrease the proton gradient generated in oxidative phosphorylation. They do this by increasing the permeability of the inner mitochondrial membrane, allowing protons that have been pumped into the intermembrane space to return to the mitochondrial matrix. UCP1-mediated heat generation in brown fat uncouples the respiratory chain, allowing for fast substrate oxidation with a low rate of ATP production. UCP1 is related to other mitochondrial metabolite transporters such as the adenine nucleotide translocator, a proton channel in the mitochondrial inner membrane that permits the translocation of protons from the mitochondrial intermembrane space to the mitochondrial matrix. UCP1 is restricted to brown adipose tissue, where it provides a mechanism for the enormous heat-generating capacity of the tissue.

UCP1 is activated in the brown fat cell by fatty acids and inhibited by nucleotides. Fatty acids cause the following signaling cascade: Sympathetic nervous system terminals release Norepinephrine onto a Beta-3 adrenergic receptor on the plasma membrane. This activates adenylyl cyclase, which catalyses the conversion of ATP to cyclic AMP (cAMP). cAMP activates protein kinase A, causing its active C subunits to be freed from its regulatory R subunits. Active protein kinase A, in turn, phosphorylates triacylglycerol lipase, thereby activating it. The lipase converts triacylglycerols into free fatty acids, which activate UCP1, overriding the inhibition caused by purine nucleotides (GDP and ADP). At the termination of thermogenesis, the mitochondria oxidize away the residual fatty acids, UCP1 inactivates and the cell resumes its normal energy-conserving mode.

History

Uncoupling protein 1 was discovered in 1978[2] and was first cloned in 1988.[3][4]

Uncoupling protein two (UCP2), a homolog of UCP1, was identified in 1997. UCP2 localizes to a wide variety of tissues, and is thought to be involved in regulating reactive oxygen species (ROS). In the past decade, three additional homologs of UCP1 have been identified, including UCP3, UCP4, and BMCP1 (also known as UCP5).

Clinical relevance

Methods of delivering UCP1 to cells by gene transfer therapy or methods of its upregulation have been an important line of enquiry in research into the treatment of obesity, due to their ability to dissipate excess metabolic stores.[5]

See also

References

  1. ↑ "Entrez Gene: UCP1 uncoupling protein 1 (mitochondrial, proton carrier)".
  2. ↑ Nicholls DG, Bernson VS, Heaton GM (1978). "The identification of the component in the inner membrane of brown adipose tissue mitochondria responsible for regulating energy dissipation". Experientia. Supplementum. 32: 89–93. doi:10.1007/978-3-0348-5559-4_9. PMID 348493.
  3. ↑ Kozak LP, Britton JH, Kozak UC, Wells JM (Sep 1988). "The mitochondrial uncoupling protein gene. Correlation of exon structure to transmembrane domains". The Journal of Biological Chemistry. 263 (25): 12274–7. PMID 3410843.
  4. ↑ Bouillaud F, Raimbault S, Ricquier D (Dec 1988). "The gene for rat uncoupling protein: complete sequence, structure of primary transcript and evolutionary relationship between exons". Biochemical and Biophysical Research Communications. 157 (2): 783–92. doi:10.1016/S0006-291X(88)80318-8. PMID 3202878.
  5. ↑ Kozak LP, Anunciado-Koza R (Dec 2008). "UCP1: its involvement and utility in obesity". International Journal of Obesity. 32 Suppl 7 (Suppl 7): S32–8. doi:10.1038/ijo.2008.236. PMC 2746324. PMID 19136989.

Further reading

  • Ricquier D, Bouillaud F (Jan 2000). "The uncoupling protein homologues: UCP1, UCP2, UCP3, StUCP and AtUCP". The Biochemical Journal. 345 Pt 2 (2): 161–79. doi:10.1042/0264-6021:3450161. PMC 1220743. PMID 10620491.
  • Muzzin P (Apr 2002). "The uncoupling proteins". Annales d'Endocrinologie. 63 (2 Pt 1): 106–10. PMID 11994670.
  • Del Mar Gonzalez-Barroso M, Ricquier D, Cassard-Doulcier AM (Oct 2000). "The human uncoupling protein-1 gene (UCP1): present status and perspectives in obesity research". Obesity Reviews. 1 (2): 61–72. doi:10.1046/j.1467-789x.2000.00009.x. PMID 12119988.
  • Cassard AM, Bouillaud F, Mattei MG, Hentz E, Raimbault S, Thomas M, Ricquier D (Jul 1990). "Human uncoupling protein gene: structure, comparison with rat gene, and assignment to the long arm of chromosome 4". Journal of Cellular Biochemistry. 43 (3): 255–64. doi:10.1002/jcb.240430306. PMID 2380264.
  • Bouillaud F, Villarroya F, Hentz E, Raimbault S, Cassard AM, Ricquier D (Jul 1988). "Detection of brown adipose tissue uncoupling protein mRNA in adult patients by a human genomic probe". Clinical Science. 75 (1): 21–7. doi:10.1042/cs0750021. PMID 3165741.
  • Oppert JM, Vohl MC, Chagnon M, Dionne FT, Cassard-Doulcier AM, Ricquier D, Pérusse L, Bouchard C (Aug 1994). "DNA polymorphism in the uncoupling protein (UCP) gene and human body fat". International Journal of Obesity and Related Metabolic Disorders. 18 (8): 526–31. PMID 7951471.
  • Clément K, Ruiz J, Cassard-Doulcier AM, Bouillaud F, Ricquier D, Basdevant A, Guy-Grand B, Froguel P (Dec 1996). "Additive effect of A-->G (-3826) variant of the uncoupling protein gene and the Trp64Arg mutation of the beta 3-adrenergic receptor gene on weight gain in morbid obesity". International Journal of Obesity and Related Metabolic Disorders. 20 (12): 1062–6. PMID 8968850.
  • Schleiff E, Shore GC, Goping IS (Mar 1997). "Human mitochondrial import receptor, Tom20p. Use of glutathione to reveal specific interactions between Tom20-glutathione S-transferase and mitochondrial precursor proteins". FEBS Letters. 404 (2–3): 314–8. doi:10.1016/S0014-5793(97)00145-2. PMID 9119086.
  • Urhammer SA, Fridberg M, Sørensen TI, Echwald SM, Andersen T, Tybjaerg-Hansen A, Clausen JO, Pedersen O (Dec 1997). "Studies of genetic variability of the uncoupling protein 1 gene in Caucasian subjects with juvenile-onset obesity". The Journal of Clinical Endocrinology and Metabolism. 82 (12): 4069–74. doi:10.1210/jc.82.12.4069. PMID 9398715.
  • Jezek P, Urbánková E (Jan 2000). "Specific sequence of motifs of mitochondrial uncoupling proteins". IUBMB Life. 49 (1): 63–70. doi:10.1080/713803586. PMID 10772343.
  • Mori H, Okazawa H, Iwamoto K, Maeda E, Hashiramoto M, Kasuga M (Mar 2001). "A polymorphism in the 5' untranslated region and a Met229-->Leu variant in exon 5 of the human UCP1 gene are associated with susceptibility to type II diabetes mellitus". Diabetologia. 44 (3): 373–6. doi:10.1007/s001250051629. PMID 11317671.
  • Nibbelink M, Moulin K, Arnaud E, Duval C, Pénicaud L, Casteilla L (Dec 2001). "Brown fat UCP1 is specifically expressed in uterine longitudinal smooth muscle cells". The Journal of Biological Chemistry. 276 (50): 47291–5. doi:10.1074/jbc.M105658200. PMID 11572862.
  • Echtay KS, Roussel D, St-Pierre J, Jekabsons MB, Cadenas S, Stuart JA, Harper JA, Roebuck SJ, Morrison A, Pickering S, Clapham JC, Brand MD (Jan 2002). "Superoxide activates mitochondrial uncoupling proteins". Nature. 415 (6867): 96–9. doi:10.1038/415096a. PMID 11780125.
  • Rousset S, del Mar Gonzalez-Barroso M, Gelly C, Pecqueur C, Bouillaud F, Ricquier D, Cassard-Doulcier AM (May 2002). "A new polymorphic site located in the human UCP1 gene controls the in vitro binding of CREB-like factor". International Journal of Obesity and Related Metabolic Disorders. 26 (5): 735–8. doi:10.1038/sj.ijo.0801973. PMID 12032762.
  • Rim JS, Kozak LP (Sep 2002). "Regulatory motifs for CREB-binding protein and Nfe2l2 transcription factors in the upstream enhancer of the mitochondrial uncoupling protein 1 gene". The Journal of Biological Chemistry. 277 (37): 34589–600. doi:10.1074/jbc.M108866200. PMID 12084707.
  • Kieć-Wilk B, Wybrańska I, Malczewska-Malec M, Leszczyńska-Gołabek L, Partyka L, Niedbał S, Jabrocka A, Dembińska-Kieć A (Sep 2002). "Correlation of the -3826A >G polymorphism in the promoter of the uncoupling protein 1 gene with obesity and metabolic disorders in obese families from southern Poland". Journal of Physiology and Pharmacology. 53 (3): 477–90. PMID 12375583.

External links

Retrieved from "https://www.wikidoc.org/index.php?title=Thermogenin&oldid=1413612"