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Currently, six homologs have been identified (UCP1-6) in mammals, with ubiquitous tissue distribution and multiple physiological functions. UCPs are regulators of key events for cellular bioenergetic metabolism, such as membrane potential, metabolic efficiency, and energy dissipation also functioning as pivotal modulators of ROS production and general cellular redox state. UCPs can act as proton channels, leading to proton re-entry the mitochondrial matrix from the intermembrane space and thus collapsing the proton gradient and decreasing the membrane potential. Each homolog exhibits its specific functions, from thermogenesis to regulation of ROS production. The expression and function of UCPs are intimately linked to diabesity, with their dysregulation\/dysfunction not only associated to diabesity onset, but also by exacerbating oxidative stress-related damage. Male infertility is one of the most overlooked diabesity-related comorbidities, where high oxidative stress takes a major role. In this review, we discuss in detail the expression and function of the different UCP homologs. In addition, the role of UCPs as key regulators of ROS production and redox homeostasis, as well as their influence on the pathophysiology of diabesity and potential role on diabesity-induced male infertility is debated.<\/jats:p>","DOI":"10.3390\/antiox10111746","type":"journal-article","created":{"date-parts":[[2021,11,1]],"date-time":"2021-11-01T11:47:38Z","timestamp":1635767258000},"page":"1746","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":36,"title":["Mitochondrial Uncoupling Proteins (UCPs) as Key Modulators of ROS Homeostasis: A Crosstalk between Diabesity and Male Infertility?"],"prefix":"10.3390","volume":"10","author":[{"given":"Bruno S.","family":"Monteiro","sequence":"first","affiliation":[{"name":"Department of Anatomy and Unit for Multidisciplinary Research in Biomedicine (UMIB), Institute of Biomedical Sciences Abel Salazar (ICBAS), University of Porto, 4050-313 Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1423-5869","authenticated-orcid":false,"given":"La\u00eds","family":"Freire-Brito","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Unit for Multidisciplinary Research in Biomedicine (UMIB), Institute of Biomedical Sciences Abel Salazar (ICBAS), University of Porto, 4050-313 Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0546-3480","authenticated-orcid":false,"given":"David F.","family":"Carrageta","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Unit for Multidisciplinary Research in Biomedicine (UMIB), Institute of Biomedical Sciences Abel Salazar (ICBAS), University of Porto, 4050-313 Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4989-5699","authenticated-orcid":false,"given":"Pedro F.","family":"Oliveira","sequence":"additional","affiliation":[{"name":"Qu\u00edmica Org\u00e2nica, Produtos Naturais e Agroalimentares (QOPNA) & Laborat\u00f3rio Associado para a Qu\u00edmica Verde\u2014Rede de Qu\u00edmica e Tecnologia (LAQV-REQUIMTE), Department of Chemistry, University of Aveiro, 3810-193 Aveiro, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7635-783X","authenticated-orcid":false,"given":"Marco G.","family":"Alves","sequence":"additional","affiliation":[{"name":"Department of Anatomy and Unit for Multidisciplinary Research in Biomedicine (UMIB), Institute of Biomedical Sciences Abel Salazar (ICBAS), University of Porto, 4050-313 Porto, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2021,10,30]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"107843","DOI":"10.1016\/j.diabres.2019.107843","article-title":"Global and regional diabetes prevalence estimates for 2019 and projections for 2030 and 2045: Results from the International Diabetes Federation Diabetes Atlas, 9(th) edition","volume":"157","author":"Saeedi","year":"2019","journal-title":"Diabetes Res. Clin. Pract."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Nuertey, B.D., Alhassan, A.I., Nuertey, A.D., Mensah, I.A., Adongo, V., Kabutey, C., Addai, J., and Biritwum, R.B. (2017). Prevalence of obesity and overweight and its associated factors among registered pensioners in Ghana; A cross sectional studies. BMC Obes., 4.","DOI":"10.1186\/s40608-017-0162-4"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1038\/sj.ijo.0803177","article-title":"Biomarkers and potential mechanisms of obesity-induced oxidant stress in humans","volume":"30","author":"Vincent","year":"2006","journal-title":"Int. J. Obes."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"902953","DOI":"10.1155\/2014\/902953","article-title":"Bioenergetics of mammalian sperm capacitation","volume":"2014","author":"Ferramosca","year":"2014","journal-title":"Biomed. Res. Int."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Carrageta, D.F., Guerra-Carvalho, B., Sousa, M., Barros, A., Oliveira, P.F., Monteiro, M.P., and Alves, M.G. (2020). Mitochondrial Activation and Reactive Oxygen-Species Overproduction during Sperm Capacitation are Independent of Glucose Stimuli. Antioxidants, 9.","DOI":"10.3390\/antiox9080750"},{"key":"ref_6","first-page":"3","article-title":"Mitochondrial electron transport chain, ROS generation and uncoupling (Review)","volume":"44","author":"Zhao","year":"2019","journal-title":"Int. J. Mol. Med."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1002\/brb3.55","article-title":"Human neuronal uncoupling proteins 4 and 5 (UCP4 and UCP5): Structural properties, regulation, and physiological role in protection against oxidative stress and mitochondrial dysfunction","volume":"2","author":"Ramsden","year":"2012","journal-title":"Brain Behav."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1882","DOI":"10.1126\/science.1130481","article-title":"Cell signaling. H2O2, a necessary evil for cell signaling","volume":"312","author":"Rhee","year":"2006","journal-title":"Science"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"37391","DOI":"10.1074\/jbc.M605552200","article-title":"Mitochondrial uncoupling protein-4 regulates calcium homeostasis and sensitivity to store depletion-induced apoptosis in neural cells","volume":"281","author":"Chan","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1300","DOI":"10.1042\/bst0311300","article-title":"Mitochondrial matrix reactive oxygen species production is very sensitive to mild uncoupling","volume":"31","author":"Miwa","year":"2003","journal-title":"Biochem. Soc. Trans."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"8705","DOI":"10.1074\/jbc.M006938200","article-title":"Uncoupling protein 2, in vivo distribution, induction upon oxidative stress, and evidence for translational regulation","volume":"276","author":"Pecqueur","year":"2001","journal-title":"J. Biol. Chem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1016\/S0014-5793(99)01568-9","article-title":"Properties of the human long and short isoforms of the uncoupling protein-3 expressed in yeast cells","volume":"462","author":"Hinz","year":"1999","journal-title":"FEBS Lett."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1038\/82565","article-title":"Disruption of the uncoupling protein-2 gene in mice reveals a role in immunity and reactive oxygen species production","volume":"26","author":"Arsenijevic","year":"2000","journal-title":"Nat. Genet."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"34611","DOI":"10.1074\/jbc.273.51.34611","article-title":"BMCP1, a novel mitochondrial carrier with high expression in the central nervous system of humans and rodents, and respiration uncoupling activity in recombinant yeast","volume":"273","author":"Sanchis","year":"1998","journal-title":"J. Biol. Chem."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"326","DOI":"10.1016\/S0014-5793(98)01713-X","article-title":"UCP4, a novel brain-specific mitochondrial protein that reduces membrane potential in mammalian cells","volume":"443","author":"Mao","year":"1999","journal-title":"FEBS Lett."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1016\/0014-5793(87)80572-0","article-title":"In the uncoupling protein from brown adipose tissue the C-terminus protrudes to the c-side of the membrane as shown by tryptic cleavage","volume":"226","author":"Eckerskorn","year":"1987","journal-title":"FEBS Lett."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"3739","DOI":"10.1002\/j.1460-2075.1993.tb06051.x","article-title":"The topology of the brown adipose tissue mitochondrial uncoupling protein determined with antibodies against its antigenic sites revealed by a library of fusion proteins","volume":"12","author":"Miroux","year":"1993","journal-title":"EMBO J."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1042\/bj3450161","article-title":"The uncoupling protein homologues: UCP1, UCP2, UCP3, StUCP and AtUCP","volume":"345","author":"Ricquier","year":"2000","journal-title":"Biochem. J."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"4004","DOI":"10.1021\/bi3003378","article-title":"Toward understanding the mechanism of ion transport activity of neuronal uncoupling proteins UCP2, UCP4, and UCP5","volume":"51","author":"Hoang","year":"2012","journal-title":"Biochemistry"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1385\/NMM:8:3:389","article-title":"Mitochondrial UCP4 mediates an adaptive shift in energy metabolism and increases the resistance of neurons to metabolic and oxidative stress","volume":"8","author":"Liu","year":"2006","journal-title":"Neuromol. Med."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1518","DOI":"10.1056\/NEJMoa0808949","article-title":"Functional brown adipose tissue in healthy adults","volume":"360","author":"Virtanen","year":"2009","journal-title":"N. Engl. J. Med."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"33370","DOI":"10.1074\/jbc.M305235200","article-title":"Acquirement of brown fat cell features by human white adipocytes","volume":"278","author":"Tiraby","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1894","DOI":"10.1038\/jid.2008.20","article-title":"Expression of uncoupling proteins in human skin and skin-derived cells","volume":"128","author":"Mori","year":"2008","journal-title":"J. Investig. Dermatol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"25073","DOI":"10.1074\/jbc.M000547200","article-title":"Thermogenic responses in brown fat cells are fully UCP1-dependent. UCP2 or UCP3 do not substitute for UCP1 in adrenergically or fatty scid-induced thermogenesis","volume":"275","author":"Matthias","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1016\/j.celrep.2013.10.044","article-title":"UCP1 in brite\/beige adipose tissue mitochondria is functionally thermogenic","volume":"5","author":"Shabalina","year":"2013","journal-title":"Cell Rep."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1016\/j.bbrc.2021.06.013","article-title":"UCP1 expression in the mouse adrenal gland is not upregulated by thermogenic conditions","volume":"566","author":"Fujita","year":"2021","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1016\/S0014-5793(02)02707-2","article-title":"Opposite regulation of uncoupling protein 1 and uncoupling protein 3 in vivo in brown adipose tissue of cold-exposed rats","volume":"519","author":"Jakus","year":"2002","journal-title":"FEBS Lett."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"C365","DOI":"10.1152\/ajpcell.00122.2018","article-title":"Cold acclimation enhances UCP1 content, lipolysis, and triacylglycerol resynthesis, but not mitochondrial uncoupling and fat oxidation, in rat white adipocytes","volume":"316","author":"Jani","year":"2019","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1007\/s10735-018-9782-3","article-title":"Cellular and subcellular localization of uncoupling protein 2 in the human kidney","volume":"49","author":"Nigro","year":"2018","journal-title":"J. Mol. Histol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"22036","DOI":"10.1074\/jbc.M412136200","article-title":"A new renal mitochondrial carrier, KMCP1, is up-regulated during tubular cell regeneration and induction of antioxidant enzymes","volume":"280","author":"Haguenauer","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1020","DOI":"10.1016\/j.placenta.2012.09.010","article-title":"Localisation and characterisation of uncoupling protein-2 (UCP2) in the human preterm placenta","volume":"33","author":"Stark","year":"2012","journal-title":"Placenta"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1159\/000494479","article-title":"UCP2 Mitigates the Loss of Human Spermatozoa Motility by Promoting mROS Elimination","volume":"50","author":"Wang","year":"2018","journal-title":"Cell Physiol. Biochem."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/S0014-5793(97)00384-0","article-title":"Uncoupling protein-3: A new member of the mitochondrial carrier family with tissue-specific expression","volume":"408","author":"Boss","year":"1997","journal-title":"FEBS Lett."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1302","DOI":"10.2337\/diabetes.50.6.1302","article-title":"Increased uncoupling protein-2 levels in beta-cells are associated with impaired glucose-stimulated insulin secretion: Mechanism of action","volume":"50","author":"Chan","year":"2001","journal-title":"Diabetes"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1038\/ng0397-269","article-title":"Uncoupling protein-2: A novel gene linked to obesity and hyperinsulinemia","volume":"15","author":"Fleury","year":"1997","journal-title":"Nat. Genet."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1073\/pnas.012410699","article-title":"A significant portion of mitochondrial proton leak in intact thymocytes depends on expression of UCP2","volume":"99","author":"Krauss","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1038\/ki.2013.80","article-title":"A microRNA-30e\/mitochondrial uncoupling protein 2 axis mediates TGF-beta1-induced tubular epithelial cell extracellular matrix production and kidney fibrosis","volume":"84","author":"Jiang","year":"2013","journal-title":"Kidney Int."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"45843","DOI":"10.1074\/jbc.M306980200","article-title":"Uncoupling protein 2, but not uncoupling protein 1, is expressed in the female mouse reproductive tract","volume":"278","author":"Rousset","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.cellimm.2013.10.002","article-title":"Mitochondrial uncoupling protein 2 protects splenocytes from oxidative stress-induced apoptosis during pathogen activation","volume":"286","author":"Cao","year":"2013","journal-title":"Cell Immunol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/j.jneuroim.2007.03.013","article-title":"Induction of Ucp2 expression in brain phagocytes and neurons following murine toxoplasmosis: An essential role of IFN-gamma and an association with negative energy balance","volume":"186","author":"Arsenijevic","year":"2007","journal-title":"J. Neuroimmunol."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Motloch, L.J., Larbig, R., Gebing, T., Reda, S., Schwaiger, A., Leitner, J., Wolny, M., Eckardt, L., and Hoppe, U.C. (2016). By Regulating Mitochondrial Ca2+-Uptake UCP2 Modulates Intracellular Ca2+. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0148359"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1016\/j.bbrc.2007.06.071","article-title":"Uncoupling protein 2 protects testicular germ cells from hyperthermia-induced apoptosis","volume":"360","author":"Zhang","year":"2007","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"12057","DOI":"10.1073\/pnas.0710434105","article-title":"UCP2 is highly expressed in pancreatic alpha-cells and influences secretion and survival","volume":"105","author":"Diao","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"E433","DOI":"10.1152\/ajpendo.2000.279.2.E433","article-title":"Impact of endotoxin on UCP homolog mRNA abundance, thermoregulation, and mitochondrial proton leak kinetics","volume":"279","author":"Yu","year":"2000","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.cyto.2006.07.012","article-title":"The uncoupling protein 2 modulates the cytokine balance in innate immunity","volume":"35","author":"Rousset","year":"2006","journal-title":"Cytokine"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"5362","DOI":"10.1074\/jbc.M807523200","article-title":"In vitro evidence suggests that miR-133a-mediated regulation of uncoupling protein 2 (UCP2) is an indispensable step in myogenic differentiation","volume":"284","author":"Chen","year":"2009","journal-title":"J. Biol. Chem."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1016\/j.bbabio.2008.03.030","article-title":"Diabetes-induced up-regulation of uncoupling protein-2 results in increased mitochondrial uncoupling in kidney proximal tubular cells","volume":"1777","author":"Friederich","year":"2008","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"760","DOI":"10.1006\/bbrc.1997.6852","article-title":"Kupffer cells are a dominant site of uncoupling protein 2 expression in rat liver","volume":"235","author":"Larrouy","year":"1997","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1111\/jnc.13244","article-title":"The mitochondrial uncoupling protein-2 is a master regulator of both M1 and M2 microglial responses","volume":"135","author":"Pandolfi","year":"2015","journal-title":"J. Neurochem."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"10417","DOI":"10.1523\/JNEUROSCI.19-23-10417.1999","article-title":"Brain uncoupling protein 2: Uncoupled neuronal mitochondria predict thermal synapses in homeostatic centers","volume":"19","author":"Horvath","year":"1999","journal-title":"J. Neurosci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1161\/01.RES.0000085581.60197.4D","article-title":"Uncoupling protein-2 overexpression inhibits mitochondrial death pathway in cardiomyocytes","volume":"93","author":"Teshima","year":"2003","journal-title":"Circ. Res."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1016\/S0014-5793(99)01540-9","article-title":"UCP2 and UCP3 rise in starved rat skeletal muscle but mitochondrial proton conductance is unchanged","volume":"462","author":"Cadenas","year":"1999","journal-title":"FEBS Lett."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"4191","DOI":"10.1016\/j.febslet.2008.11.020","article-title":"Testicular aging involves mitochondrial dysfunction as well as an increase in UCP2 levels and proton leak","volume":"582","author":"Amaral","year":"2008","journal-title":"FEBS Lett."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1482","DOI":"10.2337\/diabetes.48.7.1482","article-title":"Overexpression of uncoupling protein 2 inhibits glucose-stimulated insulin secretion from rat islets","volume":"48","author":"Chan","year":"1999","journal-title":"Diabetes"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1007\/s10863-009-9201-2","article-title":"Absolute levels of transcripts for mitochondrial uncoupling proteins UCP2, UCP3, UCP4, and UCP5 show different patterns in rat and mice tissues","volume":"41","author":"Alan","year":"2009","journal-title":"J. Bioenerg. Biomembr."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"809","DOI":"10.1096\/fasebj.11.10.9271366","article-title":"A role for uncoupling protein-2 as a regulator of mitochondrial hydrogen peroxide generation","volume":"11","author":"Hirtz","year":"1997","journal-title":"FASEB J."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.bbapap.2004.05.008","article-title":"Starvation-sensitive UCP 3 protein expression in thymus and spleen mitochondria","volume":"1700","author":"Carroll","year":"2004","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Li, Y., Maedler, K., Shu, L., and Haataja, L. (2008). UCP-2 and UCP-3 proteins are differentially regulated in pancreatic beta-cells. PLoS ONE, 3.","DOI":"10.1371\/journal.pone.0001397"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1172\/JCI16653","article-title":"Increased uncoupling protein 3 content does not affect mitochondrial function in human skeletal muscle in vivo","volume":"111","author":"Hesselink","year":"2003","journal-title":"J. Clin. Investig."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1006\/bbrc.1997.6740","article-title":"UCP3: An uncoupling protein homologue expressed preferentially and abundantly in skeletal muscle and brown adipose tissue","volume":"235","author":"Solanes","year":"1997","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/j.bbabio.2015.10.011","article-title":"The expression of UCP3 directly correlates to UCP1 abundance in brown adipose tissue","volume":"1857","author":"Hilse","year":"2016","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1064","DOI":"10.1016\/j.bbabio.2011.06.002","article-title":"Absence of mitochondrial uncoupling protein 3: Effect on thymus and spleen in the fed and fasted mice","volume":"1807","author":"Kelly","year":"2011","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"O\u2019Connor, E.B., Munoz-Wolf, N., Leon, G., Lavelle, E.C., Mills, K.H.G., Walsh, P.T., and Porter, R.K. (2020). UCP3 reciprocally controls CD4+ Th17 and Treg cell differentiation. PLoS ONE, 15.","DOI":"10.1371\/journal.pone.0239713"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"3260","DOI":"10.2337\/db12-0063","article-title":"UCP3 regulates cardiac efficiency and mitochondrial coupling in high fat-fed mice but not in leptin-deficient mice","volume":"61","author":"Boudina","year":"2012","journal-title":"Diabetes"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Huang, Z., Li, J., Du, S., Chen, G., Qi, Y., Huang, L., Xiao, L., and Tong, P. (2016). Effects of UCP4 on the Proliferation and Apoptosis of Chondrocytes: Its Possible Involvement and Regulation in Osteoarthritis. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0150684"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"2309","DOI":"10.1016\/j.bbamem.2009.07.018","article-title":"Comparative analysis of uncoupling protein 4 distribution in various tissues under physiological conditions and during development","volume":"1788","author":"Smorodchenko","year":"2009","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/j.mcn.2011.03.002","article-title":"Role of mitochondrial uncoupling protein 4 in rat inner ear","volume":"47","author":"Smorodchenko","year":"2011","journal-title":"Mol. Cell Neurosci."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.heares.2004.02.002","article-title":"Localization of the mitochondrial uncoupling protein family in the rat inner ear","volume":"196","author":"Kitahara","year":"2004","journal-title":"Hear. Res."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"993","DOI":"10.1038\/184993a0","article-title":"Thermoregulatory heat production in the brain","volume":"184","author":"Donhoffer","year":"1959","journal-title":"Nature"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1042","DOI":"10.1007\/s11064-015-1790-z","article-title":"Mast Cells Release Chemokine CCL2 in Response to Parkinsonian Toxin 1-Methyl-4-Phenyl-Pyridinium (MPP(+))","volume":"41","author":"Kempuraj","year":"2016","journal-title":"Neurochem. Res."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1016\/j.neuroscience.2005.06.056","article-title":"Regulation of mitochondrial uncoupling proteins in mouse inner ear ganglion cells in response to systemic kanamycin challenge","volume":"135","author":"Kitahara","year":"2005","journal-title":"Neuroscience"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"780","DOI":"10.1097\/01.WCB.0000122743.72175.52","article-title":"Quantitative rt-PCR analysis of uncoupling protein isoforms in mouse brain cortex: Methodological optimization and comparison of expression with brown adipose tissue and skeletal muscle","volume":"24","author":"Lengacher","year":"2004","journal-title":"J. Cereb. Blood Flow Metab."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"1611","DOI":"10.1096\/fj.99-0834com","article-title":"Characterization of novel UCP5\/BMCP1 isoforms and differential regulation of UCP4 and UCP5 expression through dietary or temperature manipulation","volume":"14","author":"Yu","year":"2000","journal-title":"FASEB J."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"658","DOI":"10.1046\/j.1471-4159.2001.00604.x","article-title":"BMCP1: A mitochondrial uncoupling protein in neurons which regulates mitochondrial function and oxidant production","volume":"79","author":"Reichert","year":"2001","journal-title":"J. Neurochem."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1016\/j.neulet.2016.02.046","article-title":"Effects of central irisin administration on the uncoupling proteins in rat brain","volume":"618","author":"Erden","year":"2016","journal-title":"Neurosci. Lett."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.neuroscience.2011.09.007","article-title":"Regulated expression of neuronal SIRT1 and related genes by aging and neuronal beta2-containing nicotinic cholinergic receptors","volume":"196","author":"Huang","year":"2011","journal-title":"Neuroscience"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"16258","DOI":"10.1074\/jbc.M910179199","article-title":"Energy metabolism in uncoupling protein 3 gene knockout mice","volume":"275","author":"Grujic","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"724","DOI":"10.1016\/j.bbabio.2019.07.010","article-title":"The human uncoupling proteins 5 and 6 (UCP5\/SLC25A14 and UCP6\/SLC25A30) transport sulfur oxyanions, phosphate and dicarboxylates","volume":"1860","author":"Gorgoglione","year":"2019","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1518","DOI":"10.1016\/j.fertnstert.2014.10.020","article-title":"Male infertility testing: Reactive oxygen species and antioxidant capacity","volume":"102","author":"Ko","year":"2014","journal-title":"Fertil. Steril."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Okamatsu-Ogura, Y., Fukano, K., Tsubota, A., Uozumi, A., Terao, A., Kimura, K., and Saito, M. (2013). Thermogenic ability of uncoupling protein 1 in beige adipocytes in mice. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0084229"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"47291","DOI":"10.1074\/jbc.M105658200","article-title":"Brown fat UCP1 is specifically expressed in uterine longitudinal smooth muscle cells","volume":"276","author":"Nibbelink","year":"2001","journal-title":"J. Biol. Chem."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"15534","DOI":"10.1074\/jbc.M413315200","article-title":"Identification of a functioning mitochondrial uncoupling protein 1 in thymus","volume":"280","author":"Carroll","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1369\/jhc.6A7013.2006","article-title":"Thymus uncoupling protein 1 is exclusive to typical brown adipocytes and is not found in thymocytes","volume":"55","author":"Frontini","year":"2007","journal-title":"J. Histochem. Cytochem."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1186\/s13059-014-0560-6","article-title":"Gateways to the FANTOM5 promoter level mammalian expression atlas","volume":"16","author":"Lizio","year":"2015","journal-title":"Genome Biol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1016\/j.freeradbiomed.2004.05.026","article-title":"The powerhouse takes control of the cell; The role of mitochondria in signal transduction","volume":"37","year":"2004","journal-title":"Free Radic. Biol. Med."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"13905","DOI":"10.1016\/S0021-9258(18)66957-1","article-title":"Expression of uncoupling protein mRNA in thermogenic or weakly thermogenic brown adipose tissue. Evidence for a rapid beta-adrenoreceptor-mediated and transcriptionally regulated step during activation of thermogenesis","volume":"261","author":"Ricquier","year":"1986","journal-title":"J. Biol. Chem."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"36","DOI":"10.3389\/fphys.2015.00036","article-title":"Mitochondrial uncoupling proteins and energy metabolism","volume":"6","author":"Busiello","year":"2015","journal-title":"Front. Physiol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1003","DOI":"10.1210\/endo.136.3.7867554","article-title":"Delineation of thyroid hormone-responsive sequences within a critical enhancer in the rat uncoupling protein gene","volume":"136","author":"Rabelo","year":"1995","journal-title":"Endocrinology"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1038\/415096a","article-title":"Superoxide activates mitochondrial uncoupling proteins","volume":"415","author":"Echtay","year":"2002","journal-title":"Nature"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"5666","DOI":"10.1074\/jbc.270.10.5666","article-title":"A novel regulatory pathway of brown fat thermogenesis. Retinoic acid is a transcriptional activator of the mitochondrial uncoupling protein gene","volume":"270","author":"Alvarez","year":"1995","journal-title":"J. Biol. Chem."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"24129","DOI":"10.1074\/jbc.272.39.24129","article-title":"Uncoupling protein-3 is a mediator of thermogenesis regulated by thyroid hormone, beta3-adrenergic agonists, and leptin","volume":"272","author":"Gong","year":"1997","journal-title":"J. Biol. Chem."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1111\/j.1432-1033.1990.tb15374.x","article-title":"Infrared spectroscopic studies of detergent-solubilized uncoupling protein from brown-adipose-tissue mitochondria","volume":"188","author":"Rial","year":"1990","journal-title":"Eur. J. Biochem."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"900","DOI":"10.2337\/diab.46.5.900","article-title":"Cloning and characterization of an uncoupling protein homolog: A potential molecular mediator of human thermogenesis","volume":"46","author":"Gimeno","year":"1997","journal-title":"Diabetes"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1831","DOI":"10.1172\/JCI200319774","article-title":"Superoxide-mediated activation of uncoupling protein 2 causes pancreatic beta cell dysfunction","volume":"112","author":"Krauss","year":"2003","journal-title":"J. Clin. Investig."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1002\/hep.20047","article-title":"Uncoupling protein-2 deficiency promotes oxidant stress and delays liver regeneration in mice","volume":"39","author":"Horimoto","year":"2004","journal-title":"Hepatology"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.cmet.2005.06.002","article-title":"Physiological functions of the mitochondrial uncoupling proteins UCP2 and UCP3","volume":"2","author":"Brand","year":"2005","journal-title":"Cell Metab."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"1200","DOI":"10.1161\/01.RES.0000170075.73039.5b","article-title":"Effects of recombinant adenovirus-mediated uncoupling protein 2 overexpression on endothelial function and apoptosis","volume":"96","author":"Lee","year":"2005","journal-title":"Circ. Res."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"42307","DOI":"10.1074\/jbc.M306951200","article-title":"Bone marrow transplantation reveals the in vivo expression of the mitochondrial uncoupling protein 2 in immune and nonimmune cells during inflammation","volume":"278","author":"Rousset","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"e41","DOI":"10.1161\/ATVBAHA.117.309228","article-title":"Reactive Oxygen Species Generation and Atherosclerosis","volume":"37","author":"Nowak","year":"2017","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"1062","DOI":"10.1038\/nm903","article-title":"Uncoupling protein-2 prevents neuronal death and diminishes brain dysfunction after stroke and brain trauma","volume":"9","author":"Mattiasson","year":"2003","journal-title":"Nat. Med."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1016\/S0006-2952(02)01167-X","article-title":"Uncoupling protein 2 (UCP2) lowers alcohol sensitivity and pain threshold","volume":"64","author":"Horvath","year":"2002","journal-title":"Biochem. Pharmacol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"442","DOI":"10.1111\/j.1440-1789.2007.00815.x","article-title":"Amplified expression of uncoupling proteins in human brain ischemic lesions","volume":"27","author":"Nakase","year":"2007","journal-title":"Neuropathology"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"33470","DOI":"10.1074\/jbc.M505258200","article-title":"Uncoupling proteins 2 and 3 function in concert to augment tolerance to cardiac ischemia","volume":"280","author":"McLeod","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"9392","DOI":"10.1073\/pnas.142206299","article-title":"Uncoupling protein 2 plays an important role in nitric oxide production of lipopolysaccharide-stimulated macrophages","volume":"99","author":"Kizaki","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/S0092-8674(01)00378-6","article-title":"Uncoupling protein-2 negatively regulates insulin secretion and is a major link between obesity, beta cell dysfunction, and type 2 diabetes","volume":"105","author":"Zhang","year":"2001","journal-title":"Cell"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"960","DOI":"10.1073\/pnas.1317400111","article-title":"UCP2 transports C4 metabolites out of mitochondria, regulating glucose and glutamine oxidation","volume":"111","author":"Vozza","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1016\/j.tibs.2009.11.001","article-title":"The on-off switches of the mitochondrial uncoupling proteins","volume":"35","author":"Azzu","year":"2010","journal-title":"Trends Biochem. Sci."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"1853","DOI":"10.1007\/s00018-007-7039-5","article-title":"Glutamine stimulates translation of uncoupling protein 2mRNA","volume":"64","author":"Hurtaud","year":"2007","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"6386","DOI":"10.1073\/pnas.94.12.6386","article-title":"Induction by leptin of uncoupling protein-2 and enzymes of fatty acid oxidation","volume":"94","author":"Zhou","year":"1997","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1042\/bj20021077","article-title":"Oxidative damage and phospholipid fatty acyl composition in skeletal muscle mitochondria from mice underexpressing or overexpressing uncoupling protein 3","volume":"368","author":"Brand","year":"2002","journal-title":"Biochem. J."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/S0014-5793(03)01386-3","article-title":"Production of endogenous matrix superoxide from mitochondrial complex I leads to activation of uncoupling protein 3","volume":"556","author":"Talbot","year":"2004","journal-title":"FEBS Lett."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"726","DOI":"10.2337\/diabetes.53.3.726","article-title":"Uncoupling proteins prevent glucose-induced neuronal oxidative stress and programmed cell death","volume":"53","author":"Vincent","year":"2004","journal-title":"Diabetes"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"2773","DOI":"10.1074\/jbc.M109736200","article-title":"The basal proton conductance of skeletal muscle mitochondria from transgenic mice overexpressing or lacking uncoupling protein-3","volume":"277","author":"Cadenas","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0378-1119(97)00596-9","article-title":"Uncoupling protein-3: A muscle-specific gene upregulated by leptin in ob\/ob mice","volume":"207","author":"Liu","year":"1998","journal-title":"Gene"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"222","DOI":"10.1006\/bbrc.1997.7636","article-title":"Regulation of the third member of the uncoupling protein family, UCP3, by cold and thyroid hormone","volume":"240","author":"Larkin","year":"1997","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1074\/jbc.273.1.5","article-title":"Uncoupling protein-3 expression in rodent skeletal muscle is modulated by food intake but not by changes in environmental temperature","volume":"273","author":"Boss","year":"1998","journal-title":"J. Biol. Chem."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"2665","DOI":"10.1172\/JCI119811","article-title":"Increased uncoupling protein-2 and -3 mRNA expression during fasting in obese and lean humans","volume":"100","author":"Millet","year":"1997","journal-title":"J. Clin. Investig."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"8713561","DOI":"10.1155\/2017\/8713561","article-title":"Neuroprotective Properties of Panax notoginseng Saponins via Preventing Oxidative Stress Injury in SAMP8 Mice","volume":"2017","author":"Huang","year":"2017","journal-title":"Evid Based Complement. Alternat. Med."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1111\/jnc.14297","article-title":"Parkinson\u2019s disease-related DJ-1 modulates the expression of uncoupling protein 4 against oxidative stress","volume":"145","author":"Xu","year":"2018","journal-title":"J. Neurochem."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"119614","DOI":"10.1016\/j.lfs.2021.119614","article-title":"High doses of butyrate induce a reversible body temperature drop through transient proton leak in mitochondria of brain neurons","volume":"278","author":"Xu","year":"2021","journal-title":"Life Sci."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"1705","DOI":"10.2147\/NDT.S107074","article-title":"Cortical spreading depression produces a neuroprotective effect activating mitochondrial uncoupling protein-5","volume":"12","author":"Viggiano","year":"2016","journal-title":"Neuropsychiatr. Dis. Treat."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1007\/978-1-4757-6125-2_15","article-title":"Oxygen and oxidative stress modulate the expression of uncoupling protein-5 in vitro and in vivo","volume":"540","author":"Pichiule","year":"2003","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"6043038","DOI":"10.1155\/2016\/6043038","article-title":"Hydrogen Sulfide and Cellular Redox Homeostasis","volume":"2016","author":"Xie","year":"2016","journal-title":"Oxid. Med. Cell. Longev."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1093\/toxsci\/kfs203","article-title":"Late-onset increases in oxidative stress and other tumorigenic activities and tumors with a Ha-ras mutation in the liver of adult male C3H mice gestationally exposed to arsenic","volume":"129","author":"Nohara","year":"2012","journal-title":"Toxicol. Sci."},{"key":"ref_125","doi-asserted-by":"crossref","unstructured":"Mydin, R., and Okekpa, S.I. (2019). Reactive oxygen species, cellular redox homeostasis and cancer. Homeostasis-An Integrated Vision, IntechOpen.","DOI":"10.5772\/intechopen.76096"},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1023\/A:1006873518427","article-title":"Reactive oxygen species, mitochondria, apoptosis and aging","volume":"174","author":"Papa","year":"1997","journal-title":"Mol. Cell. Biochem."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1016\/j.bbabio.2018.05.019","article-title":"Mitochondrial uncoupling, ROS generation and cardioprotection","volume":"1859","author":"Cadenas","year":"2018","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"1106","DOI":"10.1016\/j.freeradbiomed.2011.06.022","article-title":"Uncoupling proteins and the control of mitochondrial reactive oxygen species production","volume":"51","author":"Mailloux","year":"2011","journal-title":"Free Radic. Biol. Med."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1016\/j.bbabio.2010.02.035","article-title":"The regulation and turnover of mitochondrial uncoupling proteins","volume":"1797","author":"Azzu","year":"2010","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"1470","DOI":"10.1016\/j.bbabio.2010.04.008","article-title":"The role of UCP 1 in production of reactive oxygen species by mitochondria isolated from brown adipose tissue","volume":"1797","author":"Dlaskova","year":"2010","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"21961","DOI":"10.1074\/jbc.M110.122861","article-title":"Uncoupling protein 1 decreases superoxide production in brown adipose tissue mitochondria","volume":"285","author":"Oelkrug","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_132","first-page":"624","article-title":"Mitochondrial uncoupling prevents cold-induced oxidative stress: A case study using UCP1 knockout mice","volume":"217","author":"Stier","year":"2014","journal-title":"J. Exp. Biol."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1038\/nature17399","article-title":"Mitochondrial ROS regulate thermogenic energy expenditure and sulfenylation of UCP1","volume":"532","author":"Chouchani","year":"2016","journal-title":"Nature"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"13882","DOI":"10.1074\/jbc.M601387200","article-title":"UCP1 and defense against oxidative stress. 4-Hydroxy-2-nonenal effects on brown fat mitochondria are uncoupling protein 1-independent","volume":"281","author":"Shabalina","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"2017","DOI":"10.1016\/j.bbabio.2014.04.005","article-title":"ROS production in brown adipose tissue mitochondria: The question of UCP1-dependence","volume":"1837","author":"Shabalina","year":"2014","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1016\/j.biopha.2015.08.001","article-title":"UCP2 protects against amyloid beta toxicity and oxidative stress in primary neuronal culture","volume":"74","author":"Jun","year":"2015","journal-title":"Biomed. Pharmacother."},{"key":"ref_137","doi-asserted-by":"crossref","unstructured":"Liu, Y., Chen, L., Xu, X., Vicaut, E., and Sercombe, R. (2009). Both ischemic preconditioning and ghrelin administration protect hippocampus from ischemia\/reperfusion and upregulate uncoupling protein-2. BMC Physiol., 9.","DOI":"10.1186\/1472-6793-9-17"},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"886","DOI":"10.1161\/CIRCRESAHA.109.199018","article-title":"Transcriptional upregulation of mitochondrial uncoupling protein 2 protects against oxidative stress-associated neurogenic hypertension","volume":"105","author":"Chan","year":"2009","journal-title":"Circ. Res."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1677\/JOE-09-0117","article-title":"Uncoupling protein 2 regulates reactive oxygen species formation in islets and influences susceptibility to diabetogenic action of streptozotocin","volume":"203","author":"Lee","year":"2009","journal-title":"J. Endocrinol."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"2710","DOI":"10.2337\/db11-0132","article-title":"Beta-cell uncoupling protein 2 regulates reactive oxygen species production, which influences both insulin and glucagon secretion","volume":"60","author":"Sultan","year":"2011","journal-title":"Diabetes"},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"3040","DOI":"10.1210\/en.2008-1642","article-title":"Persistent oxidative stress due to absence of uncoupling protein 2 associated with impaired pancreatic beta-cell function","volume":"150","author":"Pi","year":"2009","journal-title":"Endocrinology"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"388","DOI":"10.1161\/01.CIR.0000051722.66074.60","article-title":"Protective role of uncoupling protein 2 in atherosclerosis","volume":"107","author":"Blanc","year":"2003","journal-title":"Circulation"},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"2813","DOI":"10.1158\/0008-5472.CAN-08-0053","article-title":"The mitochondrial uncoupling protein-2 promotes chemoresistance in cancer cells","volume":"68","author":"Derdak","year":"2008","journal-title":"Cancer Res."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1016\/j.mito.2009.12.143","article-title":"Uncoupling protein-2 and cancer","volume":"10","author":"Baffy","year":"2010","journal-title":"Mitochondrion"},{"key":"ref_145","doi-asserted-by":"crossref","unstructured":"Mailloux, R.J., Adjeitey, C.N., and Harper, M.E. (2010). Genipin-induced inhibition of uncoupling protein-2 sensitizes drug-resistant cancer cells to cytotoxic agents. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0013289"},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"31257","DOI":"10.1074\/jbc.M706129200","article-title":"Induction of endogenous uncoupling protein 3 suppresses mitochondrial oxidant emission during fatty acid-supported respiration","volume":"282","author":"Anderson","year":"2007","journal-title":"J. Biol. Chem."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1016\/j.bbabio.2005.07.001","article-title":"Uncoupling protein 3 protects aconitase against inactivation in isolated skeletal muscle mitochondria","volume":"1709","author":"Talbot","year":"2005","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"47129","DOI":"10.1074\/jbc.M208262200","article-title":"Superoxide activates mitochondrial uncoupling protein 2 from the matrix side. Studies using targeted antioxidants","volume":"277","author":"Echtay","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1002\/biof.5520240114","article-title":"Hydroxynonenal and uncoupling proteins: A model for protection against oxidative damage","volume":"24","author":"Echtay","year":"2005","journal-title":"Biofactors"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"48534","DOI":"10.1074\/jbc.M308529200","article-title":"Superoxide activates uncoupling proteins by generating carbon-centered radicals and initiating lipid peroxidation: Studies using a mitochondria-targeted spin trap derived from alpha-phenyl-N-tert-butylnitrone","volume":"278","author":"Murphy","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1016\/j.freeradbiomed.2010.05.010","article-title":"Uncoupling protein-3 lowers reactive oxygen species production in isolated mitochondria","volume":"49","author":"Toime","year":"2010","journal-title":"Free Radic. Biol. Med."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"H1192","DOI":"10.1152\/ajpheart.00592.2012","article-title":"Role of uncoupling protein 3 in ischemia-reperfusion injury, arrhythmias, and preconditioning","volume":"304","author":"Ozcan","year":"2013","journal-title":"Am. J. Physiol. Heart Circ. Physiol."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"2343","DOI":"10.2337\/diabetes.54.8.2343","article-title":"Physiological increases in uncoupling protein 3 augment fatty acid oxidation and decrease reactive oxygen species production without uncoupling respiration in muscle cells","volume":"54","author":"MacLellan","year":"2005","journal-title":"Diabetes"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.freeradbiomed.2008.09.026","article-title":"Upregulation of uncoupling protein-3 in skeletal muscle during exercise: A potential antioxidant function","volume":"46","author":"Jiang","year":"2009","journal-title":"Free Radic. Biol. Med."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"4147","DOI":"10.1016\/j.febslet.2008.11.016","article-title":"The effect of UCP3 overexpression on mitochondrial ROS production in skeletal muscle of young versus aged mice","volume":"582","author":"Nabben","year":"2008","journal-title":"FEBS Lett."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"1585","DOI":"10.1096\/fj.03-0159hyp","article-title":"A function for novel uncoupling proteins: Antioxidant defense of mitochondrial matrix by translocating fatty acid peroxides from the inner to the outer membrane leaflet","volume":"17","author":"Goglia","year":"2003","journal-title":"FASEB J."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"1148","DOI":"10.1111\/obr.12863","article-title":"Obesity and male hypogonadism: Tales of a vicious cycle","volume":"20","author":"Carrageta","year":"2019","journal-title":"Obes. Rev."},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1038\/nrendo.2017.132","article-title":"Targeting thermogenesis in brown fat and muscle to treat obesity and metabolic disease","volume":"14","author":"Betz","year":"2018","journal-title":"Nat. Rev. Endocrinol."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.jnim.2016.07.003","article-title":"Compare the resting metabolic rate status in the healthy metabolically obese with the unhealthy metabolically obese participants","volume":"6","author":"Hosseini","year":"2016","journal-title":"J. Nutr. Intermed. Metab."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"C1380","DOI":"10.1152\/ajpcell.1996.271.4.C1380","article-title":"Contribution of mitochondrial proton leak to skeletal muscle respiration and to standard metabolic rate","volume":"271","author":"Rolfe","year":"1996","journal-title":"Am. J. Physiol."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1146\/annurev.nutr.28.061807.155357","article-title":"The efficiency of cellular energy transduction and its implications for obesity","volume":"28","author":"Harper","year":"2008","journal-title":"Annu. Rev. Nutr."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"454","DOI":"10.1016\/j.cmet.2017.08.018","article-title":"The Liver as a Hub in Thermogenesis","volume":"26","author":"Abumrad","year":"2017","journal-title":"Cell Metab."},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1038\/387090a0","article-title":"Mice lacking mitochondrial uncoupling protein are cold-sensitive but not obese","volume":"387","author":"Enerback","year":"1997","journal-title":"Nature"},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1172\/JCI200315737","article-title":"Paradoxical resistance to diet-induced obesity in UCP1-deficient mice","volume":"111","author":"Liu","year":"2003","journal-title":"J. Clin. Investig."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1111\/j.1474-9726.2005.00157.x","article-title":"UCP1 deficiency increases susceptibility to diet-induced obesity with age","volume":"4","author":"Kontani","year":"2005","journal-title":"Aging Cell"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.cmet.2008.12.014","article-title":"UCP1 ablation induces obesity and abolishes diet-induced thermogenesis in mice exempt from thermal stress by living at thermoneutrality","volume":"9","author":"Feldmann","year":"2009","journal-title":"Cell Metab."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"1430","DOI":"10.1002\/oby.21542","article-title":"Sarcolipin and uncoupling protein 1 play distinct roles in diet-induced thermogenesis and do not compensate for one another","volume":"24","author":"Rowland","year":"2016","journal-title":"Obesity"},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"E515","DOI":"10.1152\/ajpendo.00097.2017","article-title":"Adaptive facultative diet-induced thermogenesis in wild-type but not in UCP1-ablated mice","volume":"313","author":"Lindsund","year":"2017","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"E729","DOI":"10.1152\/ajpendo.00020.2019","article-title":"In the absence of UCP1-mediated diet-induced thermogenesis, obesity is augmented even in the obesity-resistant 129S mouse strain","volume":"316","author":"Luijten","year":"2019","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"2366","DOI":"10.1073\/pnas.0610416104","article-title":"Ectopic brown adipose tissue in muscle provides a mechanism for differences in risk of metabolic syndrome in mice","volume":"104","author":"Almind","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"R74","DOI":"10.1152\/ajpregu.00425.2016","article-title":"Loss of UCP1 exacerbates Western diet-induced glycemic dysregulation independent of changes in body weight in female mice","volume":"312","author":"Winn","year":"2017","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"4089","DOI":"10.2337\/db14-0746","article-title":"Brown adipose tissue improves whole-body glucose homeostasis and insulin sensitivity in humans","volume":"63","author":"Chondronikola","year":"2014","journal-title":"Diabetes"},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.1056\/NEJMoa0810780","article-title":"Identification and importance of brown adipose tissue in adult humans","volume":"360","author":"Cypess","year":"2009","journal-title":"N. Engl. J. Med."},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"1385","DOI":"10.2337\/diabetes.54.5.1385","article-title":"Uncoupling protein 1 is necessary for norepinephrine-induced glucose utilization in brown adipose tissue","volume":"54","author":"Inokuma","year":"2005","journal-title":"Diabetes"},{"key":"ref_175","doi-asserted-by":"crossref","first-page":"202","DOI":"10.1016\/j.yjmcc.2015.05.002","article-title":"Functional brown adipose tissue limits cardiomyocyte injury and adverse remodeling in catecholamine-induced cardiomyopathy","volume":"84","author":"Thoonen","year":"2015","journal-title":"J. Mol. Cell. Cardiol."},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"222","DOI":"10.3389\/fendo.2020.00222","article-title":"Brown Adipose Tissue, Diet-Induced Thermogenesis, and Thermogenic Food Ingredients: From Mice to Men","volume":"11","author":"Saito","year":"2020","journal-title":"Front. Endocrinol."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"e12472","DOI":"10.1111\/jpi.12472","article-title":"Melatonin increases brown adipose tissue mass and function in Zucker diabetic fatty rats: Implications for obesity control","volume":"64","author":"Reiter","year":"2018","journal-title":"J. Pineal Res."},{"key":"ref_178","first-page":"E768","article-title":"Reduction of dietary obesity in aP2-Ucp transgenic mice: Physiology and adipose tissue distribution","volume":"270","author":"Kopecky","year":"1996","journal-title":"Am. J. Physiol."},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1007\/s12263-011-0260-8","article-title":"Augmenting energy expenditure by mitochondrial uncoupling: A role of AMP-activated protein kinase","volume":"7","author":"Klaus","year":"2012","journal-title":"Genes Nutr."},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"1115","DOI":"10.1038\/80450","article-title":"Skeletal muscle respiratory uncoupling prevents diet-induced obesity and insulin resistance in mice","volume":"6","author":"Li","year":"2000","journal-title":"Nat. Med."},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"E405","DOI":"10.1152\/ajpendo.00057.2013","article-title":"Mitochondrial uncoupling in skeletal muscle by UCP1 augments energy expenditure and glutathione content while mitigating ROS production","volume":"305","author":"Adjeitey","year":"2013","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"E495","DOI":"10.1152\/ajpendo.00518.2012","article-title":"Skeletal muscle uncoupling-induced longevity in mice is linked to increased substrate metabolism and induction of the endogenous antioxidant defense system","volume":"304","author":"Keipert","year":"2013","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1152\/physiolgenomics.00226.2007","article-title":"Uncoupling protein 1 expression in murine skeletal muscle increases AMPK activation, glucose turnover, and insulin sensitivity in vivo","volume":"33","author":"Neschen","year":"2008","journal-title":"Physiol. Genom."},{"key":"ref_184","doi-asserted-by":"crossref","unstructured":"Ost, M., Werner, F., Dokas, J., Klaus, S., and Voigt, A. (2014). Activation of AMPKalpha2 is not crucial for mitochondrial uncoupling-induced metabolic effects but required to maintain skeletal muscle integrity. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0094689"},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1210\/endo.142.1.7889","article-title":"Effects of adenoviral overexpression of uncoupling protein-2 and -3 on mitochondrial respiration in insulinoma cells","volume":"142","author":"Hong","year":"2001","journal-title":"Endocrinology"},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/j.cmet.2006.04.010","article-title":"Genipin inhibits UCP2-mediated proton leak and acutely reverses obesity- and high glucose-induced beta cell dysfunction in isolated pancreatic islets","volume":"3","author":"Zhang","year":"2006","journal-title":"Cell Metab."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"1153","DOI":"10.1096\/fj.06-7148com","article-title":"Inhibition of UCP2 expression reverses diet-induced diabetes mellitus by effects on both insulin secretion and action","volume":"21","author":"Araujo","year":"2007","journal-title":"FASEB J."},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1677\/joe.1.06715","article-title":"Endogenous islet uncoupling protein-2 expression and loss of glucose homeostasis in ob\/ob mice","volume":"190","author":"Saleh","year":"2006","journal-title":"J. Endocrinol."},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1111\/j.1463-1326.2010.01269.x","article-title":"Reactive oxygen species and uncoupling protein 2 in pancreatic beta-cell function","volume":"12","author":"Pi","year":"2010","journal-title":"Diabetes Obes. Metab."},{"key":"ref_190","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1042\/BJ20070954","article-title":"Uncoupling protein-2 contributes significantly to high mitochondrial proton leak in INS-1E insulinoma cells and attenuates glucose-stimulated insulin secretion","volume":"409","author":"Affourtit","year":"2008","journal-title":"Biochem. J."},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"1783","DOI":"10.2337\/db06-1601","article-title":"Reactive oxygen species as a signal in glucose-stimulated insulin secretion","volume":"56","author":"Pi","year":"2007","journal-title":"Diabetes"},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"1361","DOI":"10.1089\/ars.2017.7346","article-title":"Mitochondrial Reactive Oxygen Species and Type 1 Diabetes","volume":"29","author":"Chen","year":"2018","journal-title":"Antioxid. Redox Signal."},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/j.molmed.2011.08.003","article-title":"Mitochondrial uncoupling protein 2 (UCP2) in glucose and lipid metabolism","volume":"18","author":"Diano","year":"2012","journal-title":"Trends Mol. Med."},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1016\/j.freeradbiomed.2010.12.020","article-title":"Uncoupling protein-2 attenuates glucose-stimulated insulin secretion in INS-1E insulinoma cells by lowering mitochondrial reactive oxygen species","volume":"50","author":"Affourtit","year":"2011","journal-title":"Free Radic. Biol. Med."},{"key":"ref_195","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1038\/35019082","article-title":"Mice overexpressing human uncoupling protein-3 in skeletal muscle are hyperphagic and lean","volume":"406","author":"Clapham","year":"2000","journal-title":"Nature"},{"key":"ref_196","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1007\/s00125-003-1272-8","article-title":"Reduction of diet-induced obesity in transgenic mice overexpressing uncoupling protein 3 in skeletal muscle","volume":"47","author":"Son","year":"2004","journal-title":"Diabetologia"},{"key":"ref_197","doi-asserted-by":"crossref","first-page":"E1304","DOI":"10.1152\/ajpendo.00401.2005","article-title":"Effects of the presence, absence, and overexpression of uncoupling protein-3 on adiposity and fuel metabolism in congenic mice","volume":"290","author":"Costford","year":"2006","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_198","doi-asserted-by":"crossref","first-page":"4213","DOI":"10.1096\/fj.13-234302","article-title":"Muscle uncoupling protein 3 overexpression mimics endurance training and reduces circulating biomarkers of incomplete beta-oxidation","volume":"27","author":"Aguer","year":"2013","journal-title":"FASEB J."},{"key":"ref_199","doi-asserted-by":"crossref","first-page":"16251","DOI":"10.1074\/jbc.M910177199","article-title":"Lack of obesity and normal response to fasting and thyroid hormone in mice lacking uncoupling protein-3","volume":"275","author":"Gong","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_200","doi-asserted-by":"crossref","first-page":"1687","DOI":"10.1002\/oby.22879","article-title":"Loss of Uncoupling Protein 3 Attenuates Western Diet-Induced Obesity, Systemic Inflammation, and Insulin Resistance in Rats","volume":"28","author":"Lomax","year":"2020","journal-title":"Obesity"},{"key":"ref_201","doi-asserted-by":"crossref","first-page":"1995","DOI":"10.1172\/JCI13579","article-title":"Overexpression of uncoupling protein 3 in skeletal muscle protects against fat-induced insulin resistance","volume":"117","author":"Choi","year":"2007","journal-title":"J. Clin. Investig."},{"key":"ref_202","doi-asserted-by":"crossref","first-page":"E1018","DOI":"10.1152\/ajpendo.00779.2007","article-title":"Long-term high-fat feeding induces greater fat storage in mice lacking UCP3","volume":"295","author":"Costford","year":"2008","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_203","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1007\/s00424-010-0892-3","article-title":"Uncoupling protein 3 expression levels influence insulin sensitivity, fatty acid oxidation, and related signaling pathways","volume":"461","author":"Senese","year":"2011","journal-title":"Pflugers Arch."},{"key":"ref_204","doi-asserted-by":"crossref","first-page":"2870","DOI":"10.2337\/diabetes.50.12.2870","article-title":"Uncoupling protein 3 content is decreased in skeletal muscle of patients with type 2 diabetes","volume":"50","author":"Schrauwen","year":"2001","journal-title":"Diabetes"},{"key":"ref_205","doi-asserted-by":"crossref","first-page":"1520","DOI":"10.1210\/jc.2005-1572","article-title":"Reduced skeletal muscle uncoupling protein-3 content in prediabetic subjects and type 2 diabetic patients: Restoration by rosiglitazone treatment","volume":"91","author":"Schrauwen","year":"2006","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"ref_206","doi-asserted-by":"crossref","first-page":"594","DOI":"10.1111\/j.1463-1326.2006.00628.x","article-title":"Skeletal muscle uncoupling protein-3 restores upon intervention in the prediabetic and diabetic state: Implications for diabetes pathogenesis?","volume":"9","author":"Mensink","year":"2007","journal-title":"Diabetes Obes. Metab."},{"key":"ref_207","doi-asserted-by":"crossref","first-page":"0512","DOI":"10.1152\/ajplegacy.1943.138.3.512","article-title":"The role of oxygen in the metabolism and motility of human spermatozoa","volume":"138","author":"MacLeod","year":"1943","journal-title":"Am. J. Physiol."},{"key":"ref_208","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1093\/humupd\/dmn004","article-title":"Oxidative stress and male infertility\u2014A clinical perspective","volume":"14","author":"Tremellen","year":"2008","journal-title":"Hum. Reprod. Update"},{"key":"ref_209","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/S0090-4295(95)97088-6","article-title":"Incidence and level of seminal reactive oxygen species in normal men","volume":"45","author":"Shekarriz","year":"1995","journal-title":"Urology"},{"key":"ref_210","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.1439-0272.1994.tb00804.x","article-title":"Chemiluminescence in semen of infertile men","volume":"26","author":"Ochsendorf","year":"1994","journal-title":"Andrologia"},{"key":"ref_211","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1111\/j.1365-2605.1993.tb01177.x","article-title":"Reactive oxygen species in semen of infertile patients: Levels of superoxide dismutase- and catalase-like activities in seminal plasma and spermatozoa","volume":"16","author":"Zini","year":"1993","journal-title":"Int. J. Androl."},{"key":"ref_212","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1016\/S0015-0282(16)54855-9","article-title":"Formation of reactive oxygen species in spermatozoa of infertile patients","volume":"57","author":"Iwasaki","year":"1992","journal-title":"Fertil. Steril."},{"key":"ref_213","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1016\/j.fertnstert.2006.02.088","article-title":"Oxidative stress in an assisted reproductive techniques setting","volume":"86","author":"Agarwal","year":"2006","journal-title":"Fertil. Steril."},{"key":"ref_214","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1186\/1477-7827-12-33","article-title":"Characterizing semen parameters and their association with reactive oxygen species in infertile men","volume":"12","author":"Agarwal","year":"2014","journal-title":"Reprod. Biol. Endocrinol."},{"key":"ref_215","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1016\/S0015-0282(02)04948-8","article-title":"Role of reactive oxygen species in the pathophysiology of human reproduction","volume":"79","author":"Agarwal","year":"2003","journal-title":"Fertil. Steril."},{"key":"ref_216","doi-asserted-by":"crossref","first-page":"871","DOI":"10.1016\/j.fertnstert.2004.02.132","article-title":"Human sperm superoxide anion generation and correlation with semen quality in patients with male infertility","volume":"82","author":"Said","year":"2004","journal-title":"Fertil. Steril."},{"key":"ref_217","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1016\/j.fertnstert.2003.06.026","article-title":"Novel association between sperm reactive oxygen species production, sperm morphological defects, and the sperm deformity index","volume":"81","author":"Aziz","year":"2004","journal-title":"Fertil. Steril."},{"key":"ref_218","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1093\/humrep\/deh024","article-title":"Relationship between ROS production, apoptosis and DNA denaturation in spermatozoa from patients examined for infertility","volume":"19","author":"Moustafa","year":"2004","journal-title":"Hum. Reprod."},{"key":"ref_219","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1093\/humrep\/dei429","article-title":"Increased sperm DNA damage in patients with varicocele: Relationship with seminal oxidative stress","volume":"21","author":"Smith","year":"2006","journal-title":"Hum. Reprod."},{"key":"ref_220","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1530\/ror.0.0020048","article-title":"Reactive oxygen species and sperm physiology","volume":"2","author":"Jiang","year":"1997","journal-title":"Rev. Reprod."},{"key":"ref_221","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1007\/s10815-014-0425-7","article-title":"Contemporary evidence on the physiological role of reactive oxygen species in human sperm function","volume":"32","author":"Agarwal","year":"2015","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_222","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1016\/0891-5849(93)90006-G","article-title":"Human sperm hyperactivation and capacitation as parts of an oxidative process","volume":"14","author":"Gagnon","year":"1993","journal-title":"Free Radic. Biol. Med."},{"key":"ref_223","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1016\/S0002-9378(11)80017-7","article-title":"Prospective analysis of sperm-oocyte fusion and reactive oxygen species generation as criteria for the diagnosis of infertility","volume":"164","author":"Aitken","year":"1991","journal-title":"Am. J. Obstet. Gynecol."},{"key":"ref_224","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1089\/ars.2019.7977","article-title":"Endogenous and exogenous antioxidants as a tool to ameliorate male infertility induced by reactive oxygen species","volume":"33","author":"Dias","year":"2020","journal-title":"Antioxid. Redox Signal."},{"key":"ref_225","doi-asserted-by":"crossref","unstructured":"Lettieri, G., d\u2019Agostino, G., Mele, E., Cardito, C., Esposito, R., Cimmino, A., Giarra, A., Trifuoggi, M., Raimondo, S., and Notari, T. (2020). Discovery of the Involvement in DNA Oxidative Damage of Human Sperm Nuclear Basic Proteins of Healthy Young Men Living in Polluted Areas. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21124198"},{"key":"ref_226","doi-asserted-by":"crossref","unstructured":"Lettieri, G., Marra, F., Moriello, C., Prisco, M., Notari, T., Trifuoggi, M., Giarra, A., Bosco, L., Montano, L., and Piscopo, M. (2020). Molecular alterations in spermatozoa of a family case living in the land of fires. A first look at possible transgenerational effects of pollutants. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21186710"},{"key":"ref_227","doi-asserted-by":"crossref","unstructured":"Martin-Hidalgo, D., Bragado, M.J., Batista, A.R., Oliveira, P.F., and Alves, M.G. (2019). Antioxidants and Male Fertility: From Molecular Studies to Clinical Evidence. Antioxidants, 8.","DOI":"10.3390\/antiox8040089"},{"key":"ref_228","doi-asserted-by":"crossref","first-page":"613","DOI":"10.2164\/jandrol.106.001966","article-title":"Development of normal reference values for seminal reactive oxygen species and their correlation with leukocytes and semen parameters in a fertile population","volume":"28","author":"Athayde","year":"2007","journal-title":"J. Androl."},{"key":"ref_229","first-page":"780","article-title":"Correlation of sperm parameters with semen lipid peroxidation and total antioxidants levels in astheno- and oligoasheno-teratospermic men","volume":"15","author":"Karimi","year":"2013","journal-title":"Iran. Red Crescent Med. J."},{"key":"ref_230","doi-asserted-by":"crossref","first-page":"368","DOI":"10.1002\/j.1939-4640.1992.tb03327.x","article-title":"Reactive oxygen species and human spermatozoa. I. Effects on the motility of intact spermatozoa and on sperm axonemes","volume":"13","author":"Gagnon","year":"1992","journal-title":"J. Androl."},{"key":"ref_231","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1002\/j.1939-4640.1992.tb03328.x","article-title":"Reactive oxygen species and human spermatozoa. II. Depletion of adenosine triphosphate plays an important role in the inhibition of sperm motility","volume":"13","author":"Gagnon","year":"1992","journal-title":"J. Androl."},{"key":"ref_232","doi-asserted-by":"crossref","first-page":"1557988320939731","DOI":"10.1177\/1557988320939731","article-title":"Oxidative Stress is Associated with Reduced Sperm Motility in Normal Semen","volume":"14","author":"Kurkowska","year":"2020","journal-title":"Am. J. Men\u2019s Health"},{"key":"ref_233","doi-asserted-by":"crossref","first-page":"201","DOI":"10.3109\/14647273.2010.528823","article-title":"Clinical implications of sperm DNA damage","volume":"13","author":"Lewis","year":"2010","journal-title":"Hum. Fertil. (Camb.)"},{"key":"ref_234","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1016\/S1472-6483(10)60678-5","article-title":"Clinical relevance of sperm DNA damage in assisted reproduction","volume":"14","author":"Tarozzi","year":"2007","journal-title":"Reprod. Biomed. Online"},{"key":"ref_235","doi-asserted-by":"crossref","unstructured":"Cissen, M., Wely, M.V., Scholten, I., Mansell, S., Bruin, J.P., Mol, B.W., Braat, D., Repping, S., and Hamer, G. (2016). Measuring Sperm DNA Fragmentation and Clinical Outcomes of Medically Assisted Reproduction: A Systematic Review and Meta-Analysis. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0165125"},{"key":"ref_236","doi-asserted-by":"crossref","first-page":"145","DOI":"10.2164\/jandrol.111.013193","article-title":"Diabetes mellitus and sperm parameters","volume":"33","author":"Condorelli","year":"2012","journal-title":"J. Androl."},{"key":"ref_237","doi-asserted-by":"crossref","first-page":"R173","DOI":"10.1530\/REP-17-0018","article-title":"Obesity, energy balance and spermatogenesis","volume":"153","author":"Oliveira","year":"2017","journal-title":"Reproduction"},{"key":"ref_238","doi-asserted-by":"crossref","first-page":"1006","DOI":"10.1093\/humrep\/dev057","article-title":"Twenty-four-hour monitoring of scrotal temperature in obese men and men with a varicocele as a mirror of spermatogenic function","volume":"30","author":"Garolla","year":"2015","journal-title":"Hum. Reprod."},{"key":"ref_239","doi-asserted-by":"crossref","first-page":"1006","DOI":"10.1016\/S0015-0282(16)59600-9","article-title":"Association of scrotal hyperthermia with impaired spermatogenesis in infertile men","volume":"48","author":"Mieusset","year":"1987","journal-title":"Fertil. Steril."},{"key":"ref_240","first-page":"1","article-title":"Does weight loss improve semen quality and reproductive hormones? Results from a cohort of severely obese men","volume":"8","author":"Thulstrup","year":"2011","journal-title":"Reprod. Health"},{"key":"ref_241","doi-asserted-by":"crossref","first-page":"E1061","DOI":"10.1152\/ajpendo.00235.2020","article-title":"Diet during early life defines testicular lipid content and sperm quality in adulthood","volume":"319","author":"Videira","year":"2020","journal-title":"Am. J. Physiol.-Endocrinol. Metab."},{"key":"ref_242","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1530\/REP-19-0259","article-title":"A switch from high-fat to normal diet does not restore sperm quality but prevents metabolic syndrome","volume":"158","author":"Rato","year":"2019","journal-title":"Reproduction"},{"key":"ref_243","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1093\/humupd\/dmx033","article-title":"Diabetes-induced hyperglycemia impairs male reproductive function: A systematic review","volume":"24","author":"Maresch","year":"2018","journal-title":"Hum. Reprod. Update"},{"key":"ref_244","doi-asserted-by":"crossref","first-page":"626","DOI":"10.1016\/j.bbadis.2013.01.011","article-title":"Molecular mechanisms beyond glucose transport in diabetes-related male infertility","volume":"1832","author":"Alves","year":"2013","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_245","doi-asserted-by":"crossref","first-page":"e13617","DOI":"10.1111\/and.13617","article-title":"Obesity and male infertility: Mechanisms and management","volume":"53","author":"Leisegang","year":"2021","journal-title":"Andrologia"},{"key":"ref_246","doi-asserted-by":"crossref","first-page":"46","DOI":"10.2174\/157339908783502398","article-title":"Diabetes and the impairment of reproductive function: Possible role of mitochondria and reactive oxygen species","volume":"4","author":"Amaral","year":"2008","journal-title":"Curr. Diabetes Rev."},{"key":"ref_247","doi-asserted-by":"crossref","first-page":"1845","DOI":"10.1016\/j.bpj.2019.10.021","article-title":"Molecular Mechanisms Responsible for Pharmacological Effects of Genipin on Mitochondrial Proteins","volume":"117","author":"Kreiter","year":"2019","journal-title":"Biophys. 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