31:
264:
proton-motive force across mitochondrial membranes, activating the electron transport chain. Limiting the proton motive force through this process results in a negative feedback loop that limits ROS production. Especially, UCP2 decreases the transmembrane potential of mitochondria, thus decreasing the production of ROS. Thus, cancer cells may increase the production of UCP2 in mitochondria. This theory is supported by independent studies which show increased ROS production in both UCP2 and UCP3 knockout mice.
133:, which eventually led to the discovery of UCP1, initially known as "Uncoupling Protein". The brown tissue revealed elevated levels of mitochondria respiration and another respiration not coupled to ATP synthesis, which symbolized strong thermogenic activity. UCP1 was the protein discovered responsible for activating a proton pathway that was not coupled to ADP phosphorylation (ordinarily done through
276:
202:
Elsewhere in the body, uncoupling protein activities are known to affect the temperature in micro-environments. This is believed to affect other proteins' activity in these regions, though work is still required to determine the true consequences of uncoupling-induced temperature gradients within
61:
to the mitochondrial intermembrane space. The energy lost in dissipating the proton gradient via UCPs is not used to do biochemical work. Instead, heat is generated. This is what links UCP to thermogenesis. However, not every type of UCPs are related to thermogenesis. Although UCP2 and UCP3 are
294:
For example, UCPs alter the free calcium concentrations in the neuron. Mitochondria are a major site of calcium storage in neurons, and the storage capacity increases with potential across mitochondrial membranes. Therefore, when the uncoupling proteins reduce potential across these membranes,
263:
species are present. Among these activators are fatty acids, ROS, and certain ROS byproducts that are also reactive. Therefore, higher levels of ROS directly and indirectly cause increased activity of UCP2 and UCP3. This, in turn, increases proton leak from the mitochondria, lowering the
198:
show that these tissues do not function correctly without functioning uncoupling proteins. In fact, these studies revealed that cold-acclimation is not possible for these knockout mice, indicating that UCP1 is an essential driver of heat production in these brown adipose tissues.
291:, UCP2, UCP4, and UCP5 were shown to reside in neurons throughout the human central nervous system. These proteins play key roles in neuronal function. While many study findings remain controversial, several findings are widely accepted.
302:
As discussed above, neurons in the hippocampus experience increased concentrations of ATP in the presence of these uncoupling proteins. This leads scientists to hypothesize that UCPs improve synaptic plasticity and transmission.
299:, this implies UCPs play a role in regulating calcium concentrations in this region. Considering calcium ions play a large role in neurotransmission, scientists predict that these UCPs directly affect neurotransmission.
206:
The structure of human uncoupling protein 1 UCP1 has been solved by cryogenic-electron microscopy. The structure has the typical fold of a member of the SLC25 family. UCP1 is locked in a cytoplasmic-open state by
919:
Jones, S.A.; Gogoi, P.; Ruprecht, J.J.; King, M.S.; Lee, Y.; Zogg, T.; Pardon, E.; Chand, D.; Steel, S.; Coperman, D.M.; Cotrim, C.A.; Steyaert, J.; Crichton, P.G.; Moiseenkova-Bell, V.; Kunji, E.R.S. (2023).
63:
93:, for example, keeps the temperature of its spikes as much as 20 °C higher than the environment, spreading odor and attracting insects that fertilize the flowers. However, other substances, such as
194:
and small rodents, which provide non-shivering heat to these animals. These brown adipose tissues are essential to maintaining the body temperature of small rodents, and studies with (UCP1)-
145:
There are five UCP homologs known in mammals. While each of these performs unique functions, certain functions are performed by several of the homologs. The homologs are as follows:
105:
is also an uncoupling agent (chiefly in plants) and will decrease production of ATP and increase body temperature if taken in extreme excess. Uncoupling proteins are increased by
66:, which is also a proton channel. The two proteins thus work in parallel with one generating heat and the other generating ATP from ADP and inorganic phosphate, the last step in
243:
and ATP, actually resulting in a net increase in ATP concentration when these uncoupling proteins become coupled (i.e. the mechanism to allow proton leaking is inhibited).
1407:
1888:
46:) is a mitochondrial inner membrane protein that is a regulated proton channel or transporter. An uncoupling protein is thus capable of dissipating the
98:
1883:
1974:
1556:
267:
This process is important to human health, as high-concentrations of ROS are believed to be involved in the development of degenerative diseases.
62:
closely related to UCP1, UCP2 and UCP3 do not affect thermoregulatory abilities of vertebrates. UCPs are positioned in the same membrane as the
1642:
979:"Uncoupling protein-2 negatively regulates insulin secretion and is a major link between obesity, beta cell dysfunction, and type 2 diabetes"
650:
1451:
1400:
754:"UCP1 ablation induces obesity and abolishes diet-induced thermogenesis in mice exempt from thermal stress by living at thermoneutrality"
1742:
1677:
824:
Andrews ZB, Diano S, Horvath TL (November 2005). "Mitochondrial uncoupling proteins in the CNS: in support of function and survival".
1308:
Richard D, Clavel S, Huang Q, Sanchis D, Ricquier D (November 2001). "Uncoupling protein 2 in the brain: distribution and function".
1687:
1512:
1125:
Mailloux RJ, Harper ME (September 2011). "Uncoupling proteins and the control of mitochondrial reactive oxygen species production".
54:
2013:
1811:
1446:
666:
2180:
2008:
1672:
1393:
2003:
1789:
1747:
1346:, Ward MW (April 2000). "Mitochondrial membrane potential and neuronal glutamate excitotoxicity: mortality and millivolts".
295:
calcium ions are released to the surrounding environment in the neuron. Due to the high concentrations of mitochondria near
251:
The entire list of functions of UCP2 and UCP3 is not known. However, studies indicate that these proteins are involved in a
70:. Mitochondria respiration is coupled to ATP synthesis (ADP phosphorylation), but is regulated by UCPs. UCPs belong to the
2205:
686:"Uncoupling protein-3 is a mediator of thermogenesis regulated by thyroid hormone, beta3-adrenergic agonists, and leptin"
1593:
1535:
1425:
1025:"Uncoupling protein 2 prevents neuronal death including that occurring during seizures: a mechanism for preconditioning"
1647:
335:
Gaudry MJ, Jastroch M (March 2019). "Molecular evolution of uncoupling proteins and implications for brain function".
2200:
1821:
1816:
1620:
1486:
1546:
67:
30:
1737:
1615:
280:
50:
869:"Brain uncoupling protein 2: uncoupled neuronal mitochondria predict thermal synapses in homeostatic centers"
1784:
1571:
1261:"The molecular features of uncoupling protein 1 support a conventional mitochondrial carrier-like mechanism"
725:
Hagen T, Vidal-Puig A (February 2002). "Mitochondrial uncoupling proteins in human physiology and disease".
438:
Rousset S, Alves-Guerra MC, Mozo J, Miroux B, Cassard-Doulcier AM, Bouillaud F, Ricquier D (February 2004).
256:
1779:
1436:
224:
208:
86:
2153:
1893:
1769:
1605:
1583:
240:
90:
71:
259:(ROS). Current scientific consensus states that UCP2 and UCP3 perform proton transportation only when
1994:
1900:
1757:
1706:
1652:
933:
187:
130:
58:
227:
concentration with increased activity of UCP2. This is associated with cell degeneration, decreased
2210:
594:"The SLC25 Carrier Family: Important Transport Proteins in Mitochondrial Physiology and Pathology"
1924:
1566:
1371:
849:
519:
360:
1914:
1717:
1498:
1363:
1343:
1325:
1290:
1241:
1192:
1142:
1098:
1046:
1000:
959:
898:
841:
775:
734:
707:
646:
623:
571:
511:
461:
409:
352:
252:
219:
The effect of UCP2 and UCP3 on ATP concentrations varies depending on cell type. For example,
94:
1023:
Diano S, Matthews RT, Patrylo P, Yang L, Beal MF, Barnstable CJ, Horvath TL (November 2003).
1727:
1520:
1355:
1317:
1280:
1272:
1231:
1223:
1182:
1134:
1088:
1080:
1036:
990:
949:
941:
888:
880:
833:
765:
697:
613:
605:
561:
553:
501:
451:
399:
391:
344:
315:
1632:
1525:
106:
89:. Some plants species use the heat generated by uncoupling proteins for special purposes.
1138:
954:
937:
921:
1285:
1260:
1236:
1211:
1093:
1068:
893:
884:
868:
618:
593:
566:
541:
404:
379:
195:
110:
102:
1359:
995:
978:
670:
2194:
1416:
523:
296:
288:
236:
82:
1375:
977:
Zhang CY, Baffy G, Perret P, Krauss S, Peroni O, Grujic D, et al. (June 2001).
853:
364:
2049:
2024:
1682:
1561:
1474:
134:
17:
456:
439:
1276:
506:
489:
348:
1171:"Physiological functions of the mitochondrial uncoupling proteins UCP2 and UCP3"
922:"Structural basis of purine nucleotide inhibition of human uncoupling protein 1"
232:
191:
150:
114:
78:
1227:
1187:
1170:
867:
Horvath TL, Warden CH, Hajos M, Lombardi A, Goglia F, Diano S (December 1999).
770:
753:
609:
557:
275:
1801:
1463:
260:
702:
685:
395:
77:
Uncoupling proteins play a role in normal physiology, as in cold exposure or
1732:
239:. The larger number of mitochondria increases the combined concentration of
220:
1367:
1329:
1294:
1245:
1196:
1146:
1102:
1067:
Jastroch M, Divakaruni AS, Mookerjee S, Treberg JR, Brand MD (2010-06-14).
1050:
1004:
963:
945:
902:
845:
779:
738:
627:
575:
515:
465:
413:
356:
711:
1878:
1752:
1385:
1041:
1024:
645:(Fifth Edition, International ed.). China: Mary Finch. p. 668.
175:
169:
592:
Kunji, E.R.S.; King, M.S.; Ruprecht, J.J.; Thangaratnarajah, C. (2020).
2038:
2033:
1953:
1321:
1084:
228:
2143:
2138:
2108:
2103:
1983:
1963:
1958:
1948:
1943:
1938:
1933:
186:
The first uncoupling protein discovered, UCP1, was discovered in the
118:
47:
837:
752:
Feldmann HM, Golozoubova V, Cannon B, Nedergaard J (February 2009).
2163:
2158:
2148:
2133:
2128:
2123:
2118:
2113:
2098:
2093:
2088:
2083:
2078:
2073:
2068:
2063:
2058:
1854:
1841:
542:"The SLC25 Mitochondrial Carrier Family: Structure and Mechanism"
162:
156:
1389:
380:"Uncoupling proteins: current status and therapeutic prospects"
279:
This diagram shows the location of UCP1 with respect to the
235:
cells and UCP3 in muscle cells stimulate production of
231:
secretion, and type II diabetes. Conversely, UCP2 in
684:
Gong DW, He Y, Karas M, Reitman M (September 1997).
490:"Mitochondrial proton leaks and uncoupling proteins"
247:
Maintaining concentration of reactive oxygen species
2047:
2022:
1992:
1972:
1922:
1913:
1867:
1830:
1798:
1766:
1714:
1705:
1661:
1629:
1602:
1580:
1543:
1534:
1511:
1483:
1460:
1433:
1424:
378:Nedergaard J, Ricquier D, Kozak LP (October 2005).
81:, because the energy is used to generate heat (see
440:"The biology of mitochondrial uncoupling proteins"
1120:
1118:
1116:
1114:
1112:
1062:
1060:
819:
817:
815:
813:
811:
809:
129:Scientists observed the thermogenic activity in
1259:Crichton, P.G.; Lee, Y.; Kunji, E.R.S. (2017).
807:
805:
803:
801:
799:
797:
795:
793:
791:
789:
667:"California Poison Control System: Salicylates"
433:
431:
429:
427:
425:
423:
34:Structure of the human uncoupling protein UCP1
1401:
1212:"Uncoupling protein 2 and metabolic diseases"
1164:
1162:
1160:
1158:
1156:
1018:
1016:
1014:
8:
101:, also serve the same uncoupling function.
1919:
1889:Mitochondrial permeability transition pore
1871:
1711:
1540:
1430:
1408:
1394:
1386:
1284:
1235:
1186:
1092:
1069:"Mitochondrial proton and electron leaks"
1040:
994:
953:
892:
769:
701:
617:
565:
505:
455:
403:
99:carbonyl cyanide m-chlorophenyl hydrazone
1884:Mitochondrial membrane transport protein
274:
29:
327:
1643:Cholesterol side-chain cleavage enzyme
540:Ruprecht, J.J.; Kunji, E.R.S. (2020).
914:
912:
57:-powered pumping of protons from the
7:
1169:Brand MD, Esteves TC (August 2005).
587:
585:
535:
533:
483:
481:
479:
477:
475:
1557:Coenzyme Q – cytochrome c reductase
1139:10.1016/j.freeradbiomed.2011.06.022
1127:Free Radical Biology & Medicine
690:The Journal of Biological Chemistry
1743:Oxoglutarate dehydrogenase complex
1678:Glycerol-3-phosphate dehydrogenase
885:10.1523/JNEUROSCI.19-23-10417.1999
25:
1688:Carnitine palmitoyltransferase II
1812:Carbamoyl phosphate synthetase I
1452:Long-chain-fatty-acid—CoA ligase
1447:Carnitine palmitoyltransferase I
1310:Biochemical Society Transactions
1673:Glutamate aspartate transporter
1210:Sreedhar A, Zhao Y (May 2017).
641:Garrett RH, Grisham CM (2013).
1790:Pyruvate dehydrogenase complex
1748:Succinyl coenzyme A synthetase
255:limiting the concentration of
1:
1360:10.1016/s0166-2236(99)01534-9
996:10.1016/s0092-8674(01)00378-6
494:Biochim Biophys Acta Bioenerg
457:10.2337/diabetes.53.2007.S130
1594:Dihydroorotate dehydrogenase
1277:10.1016/j.biochi.2016.12.016
826:Nature Reviews. Neuroscience
507:10.1016/j.bbabio.2021.148428
349:10.1016/j.neulet.2018.12.027
287:By detecting the associated
182:Maintaining body temperature
1648:Steroid 11-beta-hydroxylase
873:The Journal of Neuroscience
2227:
1822:N-Acetylglutamate synthase
1817:Ornithine transcarbamylase
1621:Glycerol phosphate shuttle
1487:monoamine neurotransmitter
1228:10.1016/j.mito.2017.03.005
1188:10.1016/j.cmet.2005.06.002
771:10.1016/j.cmet.2008.12.014
610:10.1152/physiol.00009.2020
558:10.1016/j.tibs.2019.11.001
215:Role in ATP concentrations
211:in a pH-dependent manner.
2176:
1874:
1850:
1547:oxidative phosphorylation
223:experience a decrease in
68:oxidative phosphorylation
1738:Isocitrate dehydrogenase
1616:Malate-aspartate shuttle
703:10.1074/jbc.272.39.24129
396:10.1038/sj.embor.7400532
281:electron transport chain
1785:Glutamate dehydrogenase
1572:Succinate dehydrogenase
1348:Trends in Neurosciences
488:Nicholls, D.G. (2021).
257:reactive oxygen species
165:, also known as SLC25A9
159:, also known as SLC25A8
85:) instead of producing
2181:mitochondrial diseases
1780:Aspartate transaminase
1437:fatty acid degradation
1073:Essays in Biochemistry
946:10.1126/sciadv.adh4251
284:
253:negative-feedback loop
209:guanosine triphosphate
35:
1894:Mitochondrial carrier
1770:anaplerotic reactions
1606:mitochondrial shuttle
1584:pyrimidine metabolism
598:Physiology (Bethesda)
278:
221:pancreatic beta cells
188:brown adipose tissues
91:Eastern skunk cabbage
72:mitochondrial carrier
33:
27:Mitochondrial protein
2206:Cellular respiration
1901:Translocator protein
1758:Malate dehydrogenase
1653:Aldosterone synthase
1042:10.1210/en.2003-0667
337:Neuroscience Letters
271:Functions in neurons
174:UCP5, also known as
168:UCP4, also known as
149:UCP1, also known as
131:brown adipose tissue
59:mitochondrial matrix
1513:Intermembrane space
938:2023SciA....9H4251J
546:Trends Biochem. Sci
450:(suppl 1): S130-5.
18:Uncoupling proteins
1868:Other/to be sorted
1833:alcohol metabolism
1693:Uncoupling protein
1567:NADH dehydrogenase
1322:10.1042/bst0290812
1085:10.1042/bse0470053
285:
40:uncoupling protein
36:
2201:Membrane proteins
2188:
2187:
2172:
2171:
1915:Mitochondrial DNA
1909:
1908:
1863:
1862:
1718:citric acid cycle
1701:
1700:
1507:
1506:
1499:Monoamine oxidase
652:978-1-133-10879-5
95:2,4-dinitrophenol
16:(Redirected from
2218:
2052:
2027:
1997:
1977:
1927:
1920:
1872:
1835:
1805:
1773:
1728:Citrate synthase
1721:
1712:
1666:
1636:
1609:
1587:
1550:
1541:
1521:Adenylate kinase
1492:
1468:
1440:
1431:
1410:
1403:
1396:
1387:
1380:
1379:
1340:
1334:
1333:
1305:
1299:
1298:
1288:
1256:
1250:
1249:
1239:
1207:
1201:
1200:
1190:
1166:
1151:
1150:
1122:
1107:
1106:
1096:
1064:
1055:
1054:
1044:
1020:
1009:
1008:
998:
974:
968:
967:
957:
932:(22): eadh4251.
916:
907:
906:
896:
879:(23): 10417–27.
864:
858:
857:
821:
784:
783:
773:
749:
743:
742:
722:
716:
715:
705:
696:(39): 24129–32.
681:
675:
674:
669:. Archived from
663:
657:
656:
638:
632:
631:
621:
589:
580:
579:
569:
537:
528:
527:
509:
485:
470:
469:
459:
435:
418:
417:
407:
375:
369:
368:
332:
316:Uncoupling agent
74:(SLC25) family.
21:
2226:
2225:
2221:
2220:
2219:
2217:
2216:
2215:
2191:
2190:
2189:
2184:
2168:
2048:
2043:
2023:
2018:
1993:
1988:
1973:
1968:
1923:
1905:
1859:
1846:
1831:
1826:
1799:
1794:
1767:
1762:
1715:
1697:
1662:
1657:
1633:steroidogenesis
1630:
1625:
1603:
1598:
1581:
1576:
1544:
1530:
1526:Creatine kinase
1503:
1489:
1484:
1479:
1461:
1456:
1434:
1420:
1414:
1384:
1383:
1342:
1341:
1337:
1316:(Pt 6): 812–7.
1307:
1306:
1302:
1258:
1257:
1253:
1209:
1208:
1204:
1175:Cell Metabolism
1168:
1167:
1154:
1124:
1123:
1110:
1066:
1065:
1058:
1035:(11): 5014–21.
1022:
1021:
1012:
976:
975:
971:
918:
917:
910:
866:
865:
861:
838:10.1038/nrn1767
823:
822:
787:
758:Cell Metabolism
751:
750:
746:
724:
723:
719:
683:
682:
678:
665:
664:
660:
653:
640:
639:
635:
591:
590:
583:
539:
538:
531:
487:
486:
473:
437:
436:
421:
377:
376:
372:
334:
333:
329:
324:
312:
306:
273:
249:
217:
184:
143:
127:
107:thyroid hormone
28:
23:
22:
15:
12:
11:
5:
2224:
2222:
2214:
2213:
2208:
2203:
2193:
2192:
2186:
2185:
2177:
2174:
2173:
2170:
2169:
2167:
2166:
2161:
2156:
2151:
2146:
2141:
2136:
2131:
2126:
2121:
2116:
2111:
2106:
2101:
2096:
2091:
2086:
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2076:
2071:
2066:
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2055:
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2041:
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2020:
2019:
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2011:
2006:
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1970:
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1961:
1956:
1951:
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1941:
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1930:
1928:
1917:
1911:
1910:
1907:
1906:
1904:
1903:
1898:
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1891:
1881:
1875:
1869:
1865:
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1860:
1858:
1857:
1851:
1848:
1847:
1845:
1844:
1838:
1836:
1828:
1827:
1825:
1824:
1819:
1814:
1808:
1806:
1796:
1795:
1793:
1792:
1787:
1782:
1776:
1774:
1764:
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1760:
1755:
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1745:
1740:
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1730:
1724:
1722:
1709:
1703:
1702:
1699:
1698:
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1695:
1690:
1685:
1680:
1675:
1669:
1667:
1659:
1658:
1656:
1655:
1650:
1645:
1639:
1637:
1627:
1626:
1624:
1623:
1618:
1612:
1610:
1600:
1599:
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1596:
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1588:
1578:
1577:
1575:
1574:
1569:
1564:
1559:
1553:
1551:
1538:
1536:Inner membrane
1532:
1531:
1529:
1528:
1523:
1517:
1515:
1509:
1508:
1505:
1504:
1502:
1501:
1495:
1493:
1481:
1480:
1478:
1477:
1471:
1469:
1458:
1457:
1455:
1454:
1449:
1443:
1441:
1428:
1426:Outer membrane
1422:
1421:
1415:
1413:
1412:
1405:
1398:
1390:
1382:
1381:
1335:
1300:
1251:
1202:
1152:
1133:(6): 1106–15.
1108:
1056:
1010:
969:
908:
859:
832:(11): 829–40.
785:
744:
727:Minerva Medica
717:
676:
673:on 2014-08-02.
658:
651:
633:
604:(5): 302–327.
581:
552:(3): 244–258.
529:
471:
419:
390:(10): 917–21.
370:
326:
325:
323:
320:
319:
318:
311:
308:
297:axon terminals
272:
269:
248:
245:
216:
213:
183:
180:
179:
178:
172:
166:
160:
154:
142:
139:
126:
123:
111:norepinephrine
103:Salicylic acid
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2223:
2212:
2209:
2207:
2204:
2202:
2199:
2198:
2196:
2183:
2182:
2175:
2165:
2162:
2160:
2157:
2155:
2152:
2150:
2147:
2145:
2142:
2140:
2137:
2135:
2132:
2130:
2127:
2125:
2122:
2120:
2117:
2115:
2112:
2110:
2107:
2105:
2102:
2100:
2097:
2095:
2092:
2090:
2087:
2085:
2082:
2080:
2077:
2075:
2072:
2070:
2067:
2065:
2062:
2060:
2057:
2056:
2054:
2051:
2046:
2040:
2037:
2035:
2032:
2031:
2029:
2026:
2021:
2015:
2012:
2010:
2007:
2005:
2002:
2001:
1999:
1996:
1991:
1985:
1982:
1981:
1979:
1976:
1971:
1965:
1962:
1960:
1957:
1955:
1952:
1950:
1947:
1945:
1942:
1940:
1937:
1935:
1932:
1931:
1929:
1926:
1921:
1918:
1916:
1912:
1902:
1899:
1895:
1892:
1890:
1887:
1886:
1885:
1882:
1880:
1877:
1876:
1873:
1870:
1866:
1856:
1853:
1852:
1849:
1843:
1840:
1839:
1837:
1834:
1829:
1823:
1820:
1818:
1815:
1813:
1810:
1809:
1807:
1804:
1803:
1797:
1791:
1788:
1786:
1783:
1781:
1778:
1777:
1775:
1772:
1771:
1765:
1759:
1756:
1754:
1751:
1749:
1746:
1744:
1741:
1739:
1736:
1734:
1731:
1729:
1726:
1725:
1723:
1720:
1719:
1713:
1710:
1708:
1704:
1694:
1691:
1689:
1686:
1684:
1681:
1679:
1676:
1674:
1671:
1670:
1668:
1665:
1660:
1654:
1651:
1649:
1646:
1644:
1641:
1640:
1638:
1635:
1634:
1628:
1622:
1619:
1617:
1614:
1613:
1611:
1608:
1607:
1601:
1595:
1592:
1591:
1589:
1586:
1585:
1579:
1573:
1570:
1568:
1565:
1563:
1560:
1558:
1555:
1554:
1552:
1549:
1548:
1542:
1539:
1537:
1533:
1527:
1524:
1522:
1519:
1518:
1516:
1514:
1510:
1500:
1497:
1496:
1494:
1491:
1488:
1482:
1476:
1473:
1472:
1470:
1467:
1465:
1459:
1453:
1450:
1448:
1445:
1444:
1442:
1439:
1438:
1432:
1429:
1427:
1423:
1418:
1417:Mitochondrial
1411:
1406:
1404:
1399:
1397:
1392:
1391:
1388:
1377:
1373:
1369:
1365:
1361:
1357:
1354:(4): 166–74.
1353:
1349:
1345:
1339:
1336:
1331:
1327:
1323:
1319:
1315:
1311:
1304:
1301:
1296:
1292:
1287:
1282:
1278:
1274:
1270:
1266:
1262:
1255:
1252:
1247:
1243:
1238:
1233:
1229:
1225:
1221:
1217:
1216:Mitochondrion
1213:
1206:
1203:
1198:
1194:
1189:
1184:
1180:
1176:
1172:
1165:
1163:
1161:
1159:
1157:
1153:
1148:
1144:
1140:
1136:
1132:
1128:
1121:
1119:
1117:
1115:
1113:
1109:
1104:
1100:
1095:
1090:
1086:
1082:
1078:
1074:
1070:
1063:
1061:
1057:
1052:
1048:
1043:
1038:
1034:
1030:
1029:Endocrinology
1026:
1019:
1017:
1015:
1011:
1006:
1002:
997:
992:
989:(6): 745–55.
988:
984:
980:
973:
970:
965:
961:
956:
951:
947:
943:
939:
935:
931:
927:
923:
915:
913:
909:
904:
900:
895:
890:
886:
882:
878:
874:
870:
863:
860:
855:
851:
847:
843:
839:
835:
831:
827:
820:
818:
816:
814:
812:
810:
808:
806:
804:
802:
800:
798:
796:
794:
792:
790:
786:
781:
777:
772:
767:
763:
759:
755:
748:
745:
740:
736:
732:
728:
721:
718:
713:
709:
704:
699:
695:
691:
687:
680:
677:
672:
668:
662:
659:
654:
648:
644:
637:
634:
629:
625:
620:
615:
611:
607:
603:
599:
595:
588:
586:
582:
577:
573:
568:
563:
559:
555:
551:
547:
543:
536:
534:
530:
525:
521:
517:
513:
508:
503:
500:(7): 148428.
499:
495:
491:
484:
482:
480:
478:
476:
472:
467:
463:
458:
453:
449:
445:
441:
434:
432:
430:
428:
426:
424:
420:
415:
411:
406:
401:
397:
393:
389:
385:
381:
374:
371:
366:
362:
358:
354:
350:
346:
342:
338:
331:
328:
321:
317:
314:
313:
309:
307:
304:
300:
298:
292:
290:
282:
277:
270:
268:
265:
262:
258:
254:
246:
244:
242:
238:
234:
230:
226:
222:
214:
212:
210:
204:
200:
197:
196:knockout mice
193:
189:
181:
177:
173:
171:
167:
164:
161:
158:
155:
152:
148:
147:
146:
140:
138:
136:
132:
124:
122:
120:
116:
112:
108:
104:
100:
96:
92:
88:
84:
83:thermogenesis
80:
75:
73:
69:
65:
60:
56:
53:generated by
52:
49:
45:
41:
32:
19:
2178:
2025:ATP synthase
1832:
1800:
1768:
1716:
1692:
1683:ATP synthase
1663:
1631:
1604:
1582:
1562:Cytochrome c
1545:
1485:
1475:Kynureninase
1462:
1435:
1351:
1347:
1338:
1313:
1309:
1303:
1268:
1264:
1254:
1219:
1215:
1205:
1181:(2): 85–93.
1178:
1174:
1130:
1126:
1076:
1072:
1032:
1028:
986:
982:
972:
929:
925:
876:
872:
862:
829:
825:
764:(2): 203–9.
761:
757:
747:
733:(1): 41–57.
730:
726:
720:
693:
689:
679:
671:the original
661:
643:Biochemistry
642:
636:
601:
597:
549:
545:
497:
493:
447:
443:
387:
384:EMBO Reports
383:
373:
340:
336:
330:
305:
301:
293:
286:
266:
250:
237:mitochondria
218:
205:
201:
185:
144:
135:ATP Synthase
128:
76:
64:ATP synthase
43:
39:
37:
1975:Complex III
1344:Nicholls DG
1222:: 135–140.
343:: 140–145.
233:hippocampus
192:hibernators
151:thermogenin
115:epinephrine
79:hibernation
2211:Uncouplers
2195:Categories
1995:Complex IV
1802:urea cycle
1490:metabolism
1466:metabolism
1464:tryptophan
322:References
261:activation
153:or SLC25A7
141:In mammals
2179:see also
1925:Complex I
1733:Aconitase
1271:: 35–50.
1265:Biochimie
1079:: 53–67.
524:232774851
1879:Frataxin
1753:Fumarase
1419:proteins
1376:11564585
1368:10717676
1330:11709080
1295:28057583
1246:28351676
1197:16098826
1147:21762777
1103:20533900
1051:12960023
1005:11440717
964:37256948
955:10413660
903:10575039
854:14840725
846:16224498
780:19187776
739:11850613
628:32783608
576:31787485
516:33798544
466:14749278
444:Diabetes
414:16179945
365:56595077
357:30582970
310:See also
176:SLC25A14
170:SLC25A27
51:gradient
2039:MT-ATP8
2034:MT-ATP6
1954:MT-ND4L
1286:5395090
1237:5477468
1094:3122475
934:Bibcode
926:Sci Adv
894:6782406
712:9305858
619:7611780
567:7611774
405:1369193
229:insulin
203:cells.
125:History
2144:MT-TS2
2139:MT-TS1
2109:MT-TL2
2104:MT-TL1
2014:MT-CO3
2009:MT-CO2
2004:MT-CO1
1984:MT-CYB
1964:MT-ND6
1959:MT-ND5
1949:MT-ND4
1944:MT-ND3
1939:MT-ND2
1934:MT-ND1
1707:Matrix
1374:
1366:
1328:
1293:
1283:
1244:
1234:
1195:
1145:
1101:
1091:
1049:
1003:
962:
952:
901:
891:
852:
844:
778:
737:
710:
649:
626:
616:
574:
564:
522:
514:
464:
412:
402:
363:
355:
119:leptin
117:, and
48:proton
2164:MT-TY
2159:MT-TW
2154:MT-TV
2149:MT-TT
2134:MT-TR
2129:MT-TQ
2124:MT-TP
2119:MT-TN
2114:MT-TM
2099:MT-TK
2094:MT-TI
2089:MT-TH
2084:MT-TG
2079:MT-TF
2074:MT-TE
2069:MT-TD
2064:MT-TC
2059:MT-TA
1855:PMPCB
1842:ALDH2
1664:other
1372:S2CID
850:S2CID
520:S2CID
361:S2CID
2050:tRNA
1364:PMID
1326:PMID
1291:PMID
1242:PMID
1193:PMID
1143:PMID
1099:PMID
1047:PMID
1001:PMID
983:Cell
960:PMID
899:PMID
842:PMID
776:PMID
735:PMID
708:PMID
647:ISBN
624:PMID
572:PMID
512:PMID
498:1862
462:PMID
410:PMID
353:PMID
289:mRNA
163:UCP3
157:UCP2
97:and
55:NADH
1356:doi
1318:doi
1281:PMC
1273:doi
1269:134
1232:PMC
1224:doi
1183:doi
1135:doi
1089:PMC
1081:doi
1037:doi
1033:144
991:doi
987:105
950:PMC
942:doi
889:PMC
881:doi
834:doi
766:doi
698:doi
694:272
614:PMC
606:doi
562:PMC
554:doi
502:doi
452:doi
400:PMC
392:doi
345:doi
341:696
241:ADP
225:ATP
190:of
137:).
87:ATP
44:UCP
38:An
2197::
1370:.
1362:.
1352:23
1350:.
1324:.
1314:29
1312:.
1289:.
1279:.
1267:.
1263:.
1240:.
1230:.
1220:34
1218:.
1214:.
1191:.
1177:.
1173:.
1155:^
1141:.
1131:51
1129:.
1111:^
1097:.
1087:.
1077:47
1075:.
1071:.
1059:^
1045:.
1031:.
1027:.
1013:^
999:.
985:.
981:.
958:.
948:.
940:.
928:.
924:.
911:^
897:.
887:.
877:19
875:.
871:.
848:.
840:.
828:.
788:^
774:.
760:.
756:.
731:93
729:.
706:.
692:.
688:.
622:.
612:.
602:35
600:.
596:.
584:^
570:.
560:.
550:45
548:.
544:.
532:^
518:.
510:.
496:.
492:.
474:^
460:.
448:53
446:.
442:.
422:^
408:.
398:.
386:.
382:.
359:.
351:.
339:.
283:.
121:.
113:,
109:,
1409:e
1402:t
1395:v
1378:.
1358::
1332:.
1320::
1297:.
1275::
1248:.
1226::
1199:.
1185::
1179:2
1149:.
1137::
1105:.
1083::
1053:.
1039::
1007:.
993::
966:.
944::
936::
930:9
905:.
883::
856:.
836::
830:6
782:.
768::
762:9
741:.
714:.
700::
655:.
630:.
608::
578:.
556::
526:.
504::
468:.
454::
416:.
394::
388:6
367:.
347::
42:(
20:)
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