Selenium and the control of thyroid hormone metabolism

Thyroid

Volumn 15, Issue 8, 2005, Pages 841-853

  Köhrle, Josef ^a  

^a CHARITÉ UNIVERSITÄTSMEDIZIN BERLIN   (Germany)

Author keywords

[No Author keywords available]

Indexed keywords

DAILY ENERGY; DAILY PROTECTOR; ENZYME; GLUTATHIONE PEROXIDASE; GROWTH FACTOR; HORMONE; IODINE; IODOTHYRONINE DEIODINASE; LIOTHYRONINE; MESSENGER RNA; SELENIUM; SELENOCYSTINE; SELENOPROTEIN; SELENOPROTEIN P; THYROID BOOSTER; THYROID HELPER; THYROID HORMONE; TRACE ELEMENT; UNCLASSIFIED DRUG;

ADAPTATION; BRAIN; CANCER; CLINICAL TRIAL; ENDOCRINE GLAND; ENZYME REGULATION; GASTROINTESTINAL DISEASE; HOMEOSTASIS; HUMAN; INTENSIVE CARE; MULTIGENE FAMILY; NONHUMAN; NUTRITIONAL STATUS; PRIORITY JOURNAL; PROTEIN EXPRESSION; REVIEW; SIDE EFFECT; THYROID GLAND; THYROID HORMONE METABOLISM; THYROID HORMONE SYNTHESIS;

ANIMALS; BRAIN; CYTOKINES; HOMEOSTASIS; HUMANS; MODELS, BIOLOGICAL; RNA, MESSENGER; SELENIUM; SELENOPROTEINS; THYROID GLAND; THYROID HORMONES;

EID: 25144437445     PISSN: 10507256     EISSN: None     Source Type: Journal    
DOI: 10.1089/thy.2005.15.841     Document Type: Review

References (155)

1. Arthur JR, Beckett GJ 1999 Thyroid function. Br Med Bull 55:658-668.
2. Köhrle J 1999 The trace element selenium and the thyroid gland. Biochimie 81:527-533.
3. Leonard JL, Köhrle J 1996 Intracellular pathways of iodothyronine metabolism. In: Braverman LE, Utiger RD (eds) Werner and Ingbar's The Thyroid: A Fundamental and Clinical Text. Lippincott, Philadelphia, pp. 125-161.
4. Bassett JH, Harvey CB, Williams GR 2003 Mechanisms of thyroid hormone receptor specific nuclear and extra nuclear actions. Mol Cell Endocrinol 213:1-11.
5. Hess SY, Zimmermann MB 2004 The effect of micronutrient deficiencies on iodine nutrition and thyroid metabolism. Int J Vitamin Nutrition Res 74:103-115.
6. Bock A, Forchhammer K, Heider J, Baron C 1991 Selenoprotein synthesis: An expansion of the genetic code. Trends Biochem Sci 16:463-467.
7. Low SC, Berry MJ 1996 Knowing when not to stop: Selenocysteine incorporation in eukaryotes. Trends Biochem Sci 21:203-208.
8. Stadtman TC 1996 Selenocysteine. Annu Rev Biochem 65:83-100.
9. Hatfield DL, Gladyshev VN 2002 How selenium has altered our understanding of the genetic code. Mol Cell Biol 22:3565-3576.
10. Kryukov GV, Castellano S, Novoselov SV, et al. 2003 Characterization of mammalian selenoproteomes. Science 300:1439-1443.
11. Driscoll DM, Copeland PR 2003 Mechanism and regulation of selenoprotein synthesis. Annu Rev Nutr 23:17-40.
12. Behne D, Kyriakopoulos A, Meinhold H, Köhrle J 1990 Identification of type I iodothyronine 5′-deiodinase as a selenoenzyme. Biochem Biophys Res Commun 173:1143-1149.
13. Arthur JR, Nicol F, Beckett GJ 1990 Hepatic iodothyronine 5′-deiodinase. The role of selenium. Biochem J 272:537-540.
14. Berry MJ, Banu L, Larsen PR 1991 Type I iodothyronine deiodinase is a selenocysteine-containing enzyme. Nature 349:438-440.
15. Croteau W, Whittemore SK, Schneider MJ, St. Germain DL 1995 Cloning and expression of a cDNA for a mammalian type III iodothyronine deiodinase. J Biol Chem 270:16569-16575.
16. Croteau W, Davey JC, Galton VA, St. Germain DL 1996 Cloning of the mammalian type II iodothyronine deiodinase. J Clin Invest 98:405-417.
17. Salvatore D, Bartha T, Harney JW, Larsen PR 1996 Molecular biological and biochemical characterization of the human type 2 selenodeiodinase. Endocrinology 137:3308-3315.
18. Bianco AC, Salvatore D, Gereben B, Berry MJ, Larsen PR 2002 Biochemistry, cellular and molecular biology, and physiological roles of the iodothyronine selenodeiodinases. Endocr Rev 23:38-89.
19. Köhrle J 2002 Iodothyronine deiodinases. Methods Enzymol 347:125-167.
20. Köhrle J, Jakob F, Contempré B, Dumont JE 2005 Selenium, the thyroid, and the endocrine system. Endocrine Rev (in press).
21. Berry MJ, Kieffer JD, Harney JW, Larsen PR 1991 Selenocysteine confers the biochemical properties characteristic of the type I iodothyronine deiodinase. J Biol Chem 266: 14155-14158.
22. Salvatore D, Harvey JW, Larsen PR 1999 Mutation of the Secys residue 266 in human type 2 selenodeiodinase alters 75-Se incorporation without affecting its biochemical properties. Biochimie 81:535-538.
23. Buettner C, Harney JW, Larsen PR 2000 The role of selenocysteine 133 in catalysis by the human type 2 iodothyronine deiodinase. Endocrinology 141:4606-4612.
24. Kuiper GG, Klootwijk W, Visser TJ 2002 Substitution of cysteine for a conserved alanine residue in the catalytic center of type II iodothyronine deiodinase alters interaction with reducing cofactor. Endocrinology 143:1190-1198.
25. Kuiper GGJM, Klootwijk W, Visser TJ 2003 Substitution of cysteine for selenocysteine in the catalytic center of type III iodothyronine deiodinase reduces catalytic efficiency and alters substrate preference. Endocrinology 144:2505-2513.
26. Jacob C, Giles GI, Giles NM, Sies H 2003 Sulfur and selenium: The role of oxidation state in protein structure and function. Angew Chem Int Ed Engl 42:4742-4758.
27. Gromer S, Urig S, Becker K 2004 The thioredoxin system-From science to clinic. Med Res Rev 24:40-89.
28. Ursini F, Maiorino M, Brigelius-Flohé R, et al. 1995 Diversity of glutathione peroxidases. Methods Enzymol 252: 38-53.
29. Köhrle J, Brigelius-Flohe R, Bock A, Gartner R, Meyer O, Flohe L 2000 Selenium in biology: Facts and medical perspectives. Biol Chem 381:849-864.
30. Sies H, Packer L (eds) Protein Sensors and Reactive Oxygen Species. San Diego, Academic Press, 2002.
31. Gromer S, Johansson L, Bauer H, et al. 2003 Active sites of thioredoxin reductases: Why selenoproteins? Proc Natl Acad Sci 213:12618-12623.
32. Foster CB 2004 Selenoproteins and the Metabolic Features of the Archaeal Ancestor of Eukaryotes. Molecular Biology and Evolution msi007.
33. Mugesh G, du Mont WW, Wismach C, Jones PG 2002 Biomimetic studies on iodothyronine deiodinase intermediates: Modeling the reduction of selenenyl iodide by thiols. Chembiochem 3:440-447.
34. Roy G, Nethaji M, Mugesh G 2004 Biomimetic studies on anti-thyroid drugs and thyroid hormone synthesis. J Am Chem Soc 126:2712-2713.
35. St. Germain DL, Galton VA 1997 The deiodinase family of selenoproteins. Thyroid 7:655-668.
36. Köhrle J 2000 The deiodinase family-Selenoenzymes regulating thyroid hormone availability and action. Cell Mol Life Sci 57:1853-1863.
37. Buettner C, Harney JW, Larsen PR 1998 The 3′-untranslated region of human type 2 iodothyronine deiodinase mRNA contains a functional selenocysteine insertion sequence element. J Biol Chem 273:33374-33378.
38. Leonard DM, Stachelek SJ, Safran M, Farwell AP, Kowalik TF, Leonard JL 2000 Cloning, expression and functional characterization of the substrate binding subunit of rat type II iodothyronine 5′ deiodinase. J Biol Chem 275:25194-25201.
39. Montero-Pedrazuela A, Bernal J, Guadano-Ferraz A 2003 Divergent expression of type 2 deiodinase and the putative thyroxine-binding protein p29, in rat brain, suggests that they are functionally unrelated proteins. Endocrinol 144: 1045-1052.
40. Safran M, Leonard JL 1991 Comparison of the physicochemical properties of type I and type II iodothyronine 5′ deiodinase. J Biol Chem 266:3233-3238.
41. Leonard JL, Visser TJ, Leonard DM 2001 Characterization of the subunit structure of the catalytically active type I iodothyronine deiodinase. J Biol Chem 276:2600-2607.
42. Curcio-Morelli C, Gereben B, Zavacki AM, et al. 2003 In vivo dimerization of types 1, 2, and 3 iodothyronine selenodeiodinases. Endocrinol 144:937-946.
43. Leonard JL, Simpson GIC, Leonard DM 2005 Characterization of the protein dimerization domain responsible for the assembly of functional selenodeiodinases. J Biol Chem 280:11093-11100.
44. Berry MJ, Harney JW, Ohama T, Hatfield DL 1994 Selenocysteine insertion or termination: Factors affecting UGA codon fate and complementary anticodon:codon mutations. Nucleic Acids Res 22:3753-3759.
45. Nalvarte I, Damdimopoulos AE, Nystrom C, et al. 2004 Overexpression of enzymatically active human cytosolic and mitochondrial thioredoxin reductase in HEK-293 cells: Effect on cell growth and differentiation. J Biol Chem 279: 54510-54517.
46. Behne D, Hilmert H, Scheid S, Gessner H, Elger W 1988 Evidence for specific selenium target tissues and new biologically important selenoproteins. Biochim Biophys Acta 966:12-21.
47. Gross M, Oertel M, Köhrle J 1995 Differential selenium-dependent expression of type I 5′-deiodinase and glutathione peroxidase in the porcine epithelial kidney cell line LLC-PK1. Biochem J 306(Pt 3):851-856.
48. Muller C, Wingler K, Brigelius-Flohe R 2003 3′UTRs of glutatione peroxidases differentially affect selenium-dependent mRNA stability and selenocysteine incorporation efficiency. Biol Chem 384:11-18.
49. Safran M, Farwell AP, Leonard JL 1991 Evidence that type II 5′-deiodinase is not a selenoprotein. J Biol Chem 266:13477-13480.
50. Berry MJ, Kieffer JD, Larsen PR 1991 Evidence that cysteine, not selenocysteine, is in the catalytic site of type II iodothyronine deiodinase. Endocrinol 129:550-552.
51. Chanoine J-P, Alex S, Stone S, et al. 1993 Placental 5-deiodinase activity and fetal thyroid hormone economy are unaffected by selenium deficiency in the rat. Pediatric Res 34:288-292.
52. Köhrle J 2000 The deiodinase family: Selenoenzymes regulating thyroid hormone availability and action. Cell Mol Life Sci 57:1853-1863.
53. Bermano G, Nicol F, Dyer JA, et al. 1996 Selenoprotein gene expression during selenium-repletion of selenium-deficient rats. Biol Trace Elem Res 51:211-223.
54. Grundner-Culemann E, Martin GW III, Harney JW, Berry MJ 1999 Two distinct SECIS structures capable of directing selenocysteine incorporation in eukaryotes. RNA 5:625-635.
55. Low SC, Grundner-Culemann E, Harney JW, Berry MJ 2000 SECIS-SBP2 interactions dictate selenocysteine incorporation efficiency and selenoprotein hierarchy. EMBO J 19:6882-6890.
56. Campos-Barros A, Meinhold H, Walzog B, Behne D 1997 Effect of selenium and iodine deficiency on thyroid hormone concentrations in the central nervous system of the rat. Eur J Endocrinol 136:316-323.
57. Meinhold H, Campos-Barros A, Walzog B, Köhler R, Müller F, Behne D 1993 Effects of selenium and iodine deficiency on type I, type II, and type III iodothyronine deiodinases and circulating hormone levels in the rat. Exp Clin Endocrinol 100:87-93.
58. Bates JM, Spate VL, Morris JS, St. Germain DL, Galton VA 2000 Effects of selenium deficiency on tissue selenium content, deiodinase activity, and thyroid hormone economy in the rat during development. Endocrinol 141:2490-2500.
59. Hill KE, Lyons PR, Burk RF 1992 Differential regulation of rat liver selenoprotein mRNAs in selenium deficiency. Biochem Biophys Res Commun 185:260-263.
60. Wingler K, Bocher M, Flohe L, Kollmus H, Brigelius-Flohe R 1999 mRNA stability and selenocysteine insertion sequence efficiency rank gastrointestinal glutathione peroxidase high in the hierarchy of selenoproteins. Eur J Biochem 259:149-157.
61. Bermano G, Nicol F, Dyer JA, et al. 1995 Tissue-specific regulation of selenoenzyme gene expression during selenium deficiency in rats. Biochem J 311:425-430.
62. Pagmantidis V, Bermano G, Villette S, Broom I, Arthur J, Hesketh J 2005 Effects of Se-depletion on glutathione peroxidase and selenoprotein W gene expression in the colon. FEBS Lett 579:792-796.
63. Hill KE, Zhou J, McMahan WJ, et al. 2003 Deletion of selenoprotein P alters distribution of selenium in the mouse. J Biol Chem 278:13640-13646.
64. Schomburg L, Schweizer U, Holtmann B, Flohe L, Sendtner M, Köhrle J 2003 Gene disruption discloses role of selenoprotein P in selenium delivery to target tissues. Biochem J 370:397-402.
65. Schweizer U, Streckfuss F, Pelt P, et al. 2005 Hepatically-derived selenoprotein P is a key factor for kidney but not for brain selenium supply. Biochem J 386:221-226.
66. Sun Y, Butler JA, Whanger PD 2001 Glutathione peroxidase activity and selenoprotein W levels in different brain regions of selenium depleted rats. J Nutr Biochem 12:88-94.
67. Hill KE, Xia Y, Akesson B, Boeglin ME, Burk RF 1996 Selenoprotein P concentration in plasma is an index of selenium status in selenium-deficient and selenium-supplemented Chinese subjects. J Nutr 126:138-145.
68. Persson-Moschos M, Alfthan G, Akesson B 1998 Plasma selenoprotein P levels of healthy males in different selenium status after oral supplementation with different forms of selenium. Eur J Clin Nutr 52:363-367.
69. Roti E, Minelli R, Gardini E, et al. 1993 Selenium administration does not cause thyroid insufficiency in subjects with mild iodine deficiency and sufficient selenium intake. J Endocrinol Invest 16:481-484.
70. Contempré B, Dumont JE, Bebe N, Thilly CH, Diplock AT, Vanderpas J 1991 Effect of selenium supplementation in hypothyroid subjects of an iodine and selenium deficient area: The possible danger of indiscriminate supplementation of iodine deficient subjects with selenium. J Clin Endocrinol Metab 73:213-215.
71. Ravaglia G, Forti P, Maioli F, et al. 2000 Blood micronutrient and thyroid hormone concentrations in the oldest-old. J Clin Endocrinol Metab 85:2260-2265.
72. Olivieri O, Girelli D, Azzini M, et al. 1995 Low selenium status in the elderly influences thyroid hormones. Clin Sci Colch 89:637-642.
73. Olivieri O, Girelli D, Stanzial AM, Rossi L, Bassi A, Corrocher R 1996 Selenium, zinc, and thyroid hormones in healthy subjects: Low T3/T4 ratio in the elderly is related to impaired selenium status. Biol Trace Elem Res 51:31-41.
74. Derumeaux H, Valeix P, Castetbon K, et al. 2003 Association of selenium with thyroid volume and echostructure in 35- to 60-year-old French adults. Eur J Endocrinol 148:309-315.
75. Kvicala J, Zamrazil V, Soutorova M, Tomiska F 1995 Correlations between parameters of body selenium status and peripheral thyroid parameters in the low selenium region. Analyst 120:959-965.
76. Kvicala J, Zamrazil V 2003 Effect of iodine and selenium upon thyroid function. Cent Eur J Public Health 11:107-113.
77. Feldkamp J, Dohan O, Becker A, Seppel T, Szabolcz I, Lombeck I 1996 Selenium and iodide status of 301 institutionalized elderly subjects. Exp Clin Endocrinol 104:92.
78. Lassen KO, Horder M 1994 Selenium status and the effect of organic and inorganic selenium supplementation in a group of elderly people in Denmark. Scand J Clin Lab Invest 54:585-590.
79. Savarino L, Granchi D, Ciapetti G, et al. 2001 Serum concentrations of zinc and selenium in elderly people: Results in healthy nonagenarians/ centenarians. Exp Gerontol 36: 327-339.
80. Duffield AJ, Thomson CD, Hill KE, Williams S 1999 An estimation of selenium requirements for New Zealanders. Am J Clin Nutr 70:896-903.
81. Thomson CD 2003 Selenium and iodine interactions with thyroid status. Asia Pac J Clin Nutr 12(suppl):S14.
82. Hagmar L, Persson-Moschos M, Akesson B, Schutz A 1998 Plasma levels of selenium, selenoprotein P and glutathione peroxidase and their correlations to fish intake and serum levels of thyrotropin and thyroid hormones: A study on Latvian fish consumers. Eur J Clin Nutr 52:796-800.
83. Hawkes WC, Keim NL 2003 Dietary selenium intake modulates thyroid hormone and energy metabolism in men. J Nutr 133:3443-3448.
84. Hawkes WC, Alkan FZ, Oehler L 2003 Absorption, distribution and excretion of selenium from beef and rice in healthy North American men. J Nutr 133:3434-3442.
85. Brätter P, Negretti De Brätter VE 1996 Influence of high dietary selenium intake on the thyroid hormone level in human serum. J Trace Elem Med Biol 10:163-166.
86. Berger MM, Lemarchand Beraud T, Cavadini C, Chiolero R 1996 Relations between the selenium status and the low T3 syndrome after major trauma. Intens Care Med 22: 575-581.
87. Berger MM, Reymond MJ, Shenkin A, et al. 2001 Influence of selenium supplements on the post-traumatic alterations of the thyroid axis: A placebo-controlled trial. Intens Care Med 27:91-100.
88. Peeters RP, Wouters PJ, Kaptein E, van Toor H, Visser TJ, Van den Berghe G 2003 Reduced activation and increased inactivation of thyroid hormone in tissues of critically ill patients. J Clin Endocrinol Metab 88:3202-3211.
89. Mostert V 2000 Selenoprotein P: Properties, functions, and regulation. Arch Biochem Biophys 376:433-438.
90. Mostert V, Wolff S, Dreher I, Köhrle J, Abel J 2001 Identification of an element within the promoter of human selenoprotein P responsive to transforming growth factor-beta. Eur J Biochem 268:6176-6181.
91. Strain JJ, Bokje E, van't Veer P, et al. 1997 Thyroid hormones and selenium status in breast cancer. Nutr Cancer 27:48-52.
92. Makropoulos W, Heintz B, Stefanidis I 1997 Selenium deficiency and thyroid function in acute renal failure. Renal Failure 19:129-136.
93. Holzer R, Bockenkamp B, Booker P, Newland P, Ciotti G, Pozzi M 2004 The impact of cardiopulmonary bypass on selenium status, thyroid function, and oxidative defense in children. Pediatr Cardiol 25:522-528.
94. Van Lente F, Daher R 1992 Plasma selenium concentrations in patients with euthyroid sick syndrome. Clin Chem 38:1885-1888.
95. Forceville X 2001 Selenium and the "free" electron. Selenium-A trace to be followed in septic or inflammatory ICU patients? Intens Care Med 27:16-18.
96. Schilling JU, Zimmermann T, Albrecht S, Zwipp H, Saeger HD 1999 [Low T3 syndrome in multiple trauma patients-A phenomenon or important pathogenetic factor?] Med Klin 94(Suppl 3):66-69.
97. Angstwurm MW, Schopohl J, Gaertner R 2004 Selenium substitution has no direct effect on thyroid hormone metabolism in critically ill patients. Eur J Endocrinol 151: 47-54.
98. Angstwurm MWA, Schottdorf J, Schopohl J, Gärtner R 1999 Selenium replacement in patients with severe systemic inflammatory response syndrome improves clinical outcome. Crit Care Med 27:1807-1813.
99. Van den Berghe G 2000 Novel insights into the neuroendocrinology of critical illness. Eur J Endocrinol 143:1-13.
100. Dumont JE, Corvilain B, Contempre B 1994 The biochemistry of endemic cretinism: Roles of iodine and selenium deficiency and goitrogens. MoI Cell Endocrinol 100:163-166.
101. Calomme MR, Vanderpas JB, Francois B, et al. 1995 Thyroid function parameters during a selenium repletion/ depletion study in phenylketonuric subjects. Experientia 51:1208-1215.
102. Calomme M, Vanderpas J, Francois G, et al. 1995 Effects of selenium supplementation on thyroid hormone metabolism in phenylketonuria subjects on a phenylalanine restricted diet. Biol Trace Elem Res 47:349-353.
103. van Bakel MM, Printzen G, Wermuth B, Wiesmann UN 2000 Antioxidant and thyroid hormone status in selenium-deficient phenylketonuric and hyperphenylalaninemic patients. Am J Clin Nutr 72:976-981.
104. Kauf E, Dawczynski H, Jahreis G, Janitzky E, Winnefeld K 1994 Sodium selenite therapy and thyroid-hormone status in cystic fibrosis and congenital hypothyroidism. Biol Trace Elem Res 40:247-253.
105. Terwolbeck K, Behne D, Meinhold H, Menzel H, Lombeck I 1993 Increased plasma T4-levels in children with low selenium state due to reduced type I iodothyronine 5′-deiodinase activity? J Trace Elem Electrolytes Health Dis 7:53-55.
106. Chanoine JP, Neve J, Wu S, Vanderpas J, Bourdoux P 2001 Selenium decreases thyroglobulin concentrations but does not affect the increased thyroxine-totriiodothyronine ratio in children with congenital hypothyroidism. J Clin Endocrinol Metab 86:1160-1163.
107. Chanoine JP 2003 Selenium and thyroid function in infants, children and adolescents. Biofactors 19:137-143.
108. Klinger G, Shamir R, Singer P, Diamond EM, Josefsberg Z, Sirota L 1999 Parenteral selenium supplementation in extremely low birth weight infants: Inadequate dosage but no correlation with hypothyroidism. J Perinatol 19:568-572.
109. Zimmermann MB, Köhrle J 2002 The impact of iron and selenium deficiencies on iodine and thyroid metabolism: Biochemistry and relevance to public health. Thyroid 12:867-878.
110. Rayman MP 2000 The importance of selenium to human health. Lancet 356:233-241.
111. Aydin K, Kendirci M, Kurtoglu S, Karakucuk EI, Kiris A 2002 Iodine and selenium deficiency in school-children in an endemic goiter area in Turkey. J Pediatr Endocrinol Metab 15:1027-1031.
112. Zimmermann MB, Adou P, Torresani T, Zeder C, Hurrell RF 2000 Effect of oral iodized oil on thyroid size and thyroid hormone metabolism in children with concurrent selenium and iodine deficiency. Eur J Clin Nutr 54:209-213.
113. Ewan RC 1976 Effect of selenium on rat growth, growth hormone and diet utilization. J Nutr 106:702-709.
114. Thorlacius-Ussing O, Flyvbjerg A, Esmann J 1987 Evidence that selenium induces growth retardation through reduced growth hormone and somatomedin C production. Endocrinol 120:659-663.
115. Thompson KM, Haibach H, Sunde RA 1995 Growth and plasma triiodothyronine concentrations are modified by selenium deficiency and repletion in second-generation selenium-deficient rats. J Nutr 125:864-873.
116. Jorgensen JO, Moller J, Laursen T, Orskov H, Christiansen JS, Weeke J 1994 Growth hormone administration stimulates energy expenditure and extrathyroidal conversion of thyroxine to rriiodothyronine in a dose-dependent manner and suppresses circadian thyrotrophin levels: Studies in GH-deficient adults. Clin Endocrinol 41:609-614.
117. Contempré B, Duale NL, Dumont JE, Ngo B, Diplock AT, Vanderpas J 1992 Effect of selenium supplementation on thyroid hormone metabolism in an iodine and selenium deficient population. Clin Endocrinol (Oxf) 36:579-583.
118. Vanderpas JB, Contempré B, Duale NL, et al. 1993 Selenium deficiency mitigates hypothyroxinemia in iodine-deficient subjects. Am J Clin Nutr 57:2715-2755.
119. Golstein J, Corvilain B, Lamy F, Paquer D, Dumont JE 1988 Effects of a selenium deficient diet on thyroid function of normal and perchlorate treated rats. Acta Endocrinol (Copenh) 118:495-502.
120. Brabant G, Bergmann P, Kirsch CM, Köhrle J, Hesch RD, Von zur Mühlen A 1992 Alterations in the temporal pattern of thyrotropin and thyroglobulin secretion in man with experimental iodine depletion. Metabolism 41:1093-1096.
121. Contempré B, Dumont JE, Denef J-F, Many M-C 1995 Effects of selenium deficiency on thyroid necrosis, fibrosis and proliferation: A possible role in myxoedematous cretinism. Eur J Endocrinol 133:99-109.
122. Contempré B, Denef JF, Dumont JE, Many MC 1993 Selenium deficiency aggravates the necrotizing effects of a high iodide dose in iodine deficient rats. Endocrinol 132:1866-1868.
123. Contempré B, de Escobar GM, Denef JF, Dumont JE, Many MC 2004 Thiocyanate induces cell necrosis and fibrosis in selenium- and iodine-deficient rat thyroids: A potential experimental model for myxedematous endemic creteinism in Central Africa. Endocrinol 145:994-1002.
124. Contempre B, Le Moine O, Dumont JE, Denef J-F, Many MC 1996 Selenium deficiency and thyroid fibrosis. A key role for macrophages and transforming growth factor-(TGF-). Mol Cell Endocrinol 124:7-15.
125. Colzani RM, Alex S, Fang SL, Stone S, Braverman LE 1999 Effects of iodine repletion on thyroid morphology in iodine and/or selenium deficient rat term fetuses, pups and mothers. Biochimie 81:485-491.
126. Arthur JR, Nicol F, Beckett GJ 1993 Selenium deficiency, thyroid hormone metabolism, and thyroid hormone deiodinases. Am J Clin Nutr 57:2365-2395.
127. Visser TJ 1996 Pathways of thyroid hormone metabolism. Acta Medica Austriaca 23:10-16.
128. Wu SY, Huang WS, Chopra IJ, Jordan M, Alvarez D, Santini F 1995 Sulfation pathway of thyroid hormone metabolism in selenium-deficient male rats. AJP-Renal Physiol 268:E572-579.
129. Moreno-Reyes R, Suetens C, Mathieu F, et al. 1998 Kashin-Beck osteoarthropathy in rural Tibet in relation to selenium and iodine status. N Engl J Med 339:1112-11120.
130. Moreno-Reyes R, Egrise D, Neve J, Pasteels JL, Schoutens A 2001 Selenium deficiency-induced growth retardation is associated with an impaired bone metabolism and osteopenia. J Bone Miner Res 16:1556-1563.
131. Moreno-Reyes R, Mathieu F, Boelaert M, et al. 2003 Selenium and iodine supplementation of rural Tibetan children affected by Kashin-Beck osteoarthropathy. Am J Clin Nutr 78:137-144.
132. Meinhold H, Campos-Barros A, Behne D 1992 Effects of selenium and iodine deficiency on iodothyronine deiodinases in brain, thyroid and peripheral tissue. Acta Medica Austriaca 19:8-12.
133. Chanoine JP, Safran M, Farwell AP, et al. 1992 Selenium deficiency and type II 5′-deiodinase regulation in the euthryoid and hypothyroid rat: Evidence of a direct effect of thyroxine. Endocrinology 131:479-484.
134. Eder K, Kralik A, Kirchgessner M 1995 [Effect on metabolism of thyroid hormones in deficient to subtoxic selenium supply levels]. Z Ernahrungswiss 34:277-283.
135. Christensen MJ, Cammack PM, Wray CD 1995 Tissue specificity of selenoprotein gene expression in rats. J Nutr Biochem 6:367-372.
136. Beech SG, Walker SW, Beckett GJ, Arthur JR, Nicol F, Lee D 1995 Effect of selenium depletion on thyroidal type-I iodothyronine deiodinase activity in isolated human thyrocytes and rat thyroid and liver. Analyst 120:827-831.
137. Mitchell JH, Nicol F, Beckett GJ, Arthur JR 1998 Selenoprotein expression and brain development in preweanling selenium- and iodine-deficient rats. J Mol Endocrinol 20:203-210.
138. Mitchell JH, Nicol F, Beckett GJ, Arthur JR 1997 Selenium and iodine deficiencies: Effects on brain and brown adipose tissue selenozyme activity and expression. J Endocrinol 155:255-263.
139. Mitchell JH, Nicol F, Beckett GJ, Arthur JR 1996 Selenoenzyme expression in thyroid and liver of second generation selenium- and iodine-deficient rats. J Mol Endocrinol 16:259-267.
140. Bates JM, St. Germain DL, Galton VA 1999 Expression profiles of three iodothyronine deiodinases D1, D2, D3, in the developing rat. Endocrinol 140:844-851.
141. Schweizer U, Michaelis M, Köhrle J, Schomburg L 2004 Efficient selenium transfer from mother to offspring in selenoprotein-P-deficient mice enables dose-dependent rescue of phenotypes associated with selenium deficiency. Biochem J 378:21-26.
142. Hill KE, Zhou J, McMahan WJ, Motley AK, Burk RF 2004 Neurological dysfunction occurs in mice with targeted deletion of the selenoprotein P gene. J Nutr 134:157-161.
143. Cammack PM, Zwahlen BA, Christensen MJ 1995 Selenium deficiency alters thyroid hormone metabolism in guinea pigs. J Nutr 125:302-308.
144. Zagrodzki P, Nicol F, McCoy MA, et al. 1998 Iodine deficiency in cattle: Compensatory changes in thyroidal selenoenzymes. Res Vet Sci 64:209-211.
145. Reyns GE, Venken K, Morreale DE, Kuhn ER, Darras VM 2003 Dynamics and regulation of intracellular thyroid hormone concentrations in embryonic chicken liver, kidney, brain, and blood. Gen Comp Endocrinol 134:80-87.
146. Yu S, Howard KA, Wedekind KJ, Morris JG, Rogers QR 2002 A low-selenium diet increases thyroxine and decreases 3,5,3′triiodothyronine in the plasma of kittens. J Anim Physiol Anim Nutr (Berl) 86:36-41.
147. Kang BP, Bansal MP, Mehta U 2000 Hyperlipidemia and type I 5′-monodeiodinase activity: Regulation by selenium supplementation in rabbits. Biol Trace Elem Res 77:231-239.
148. Dhingra S, Singh U, Bansal MP 2004 Effect of selenium depletion and supplementation on the kinetics of type I 5′-iodothyronine deiodinase and T3/T4 in rats. Biol Trace Elem Res 97:95-104.
149. Reid ME, Stratton MS, Lillico AJ, et al. 2004 A report of high-dose selenium supplementation: Response and toxicities. J Trace Elem Med Biol 18:69-74.
150. Yang G, Zhou R 1994 Further observations on the human maximum safe dietary selenium intake in a seleniferous area of China. J Trace Elem Electrolytes Health Dis 8:159-165.
151. Bähr K, Dreher I, Köhrle J 1999 Selenium supplementation by selenium yeast and sodium selenite. Analysis of selenium state as well as risks of deficiency and intoxication. Lab Med 23:594-599.
152. Tinggi U 2003 Essentiality and toxicity of selenium and its status in Australia: A review. Toxicol Lett 137:103-110.
153. Schrauzer GN 2000 Selenomethionine: A review of its nutritional significance, metabolism and toxicity. J Nutr 130:1653-1656.
154. Combs GF, Gray WP 1998 Chemopreventive agents: Selenium. Pharmacol Ther 79:179-192.
155. Combs GF Jr 1997 Dietary selenium allowances and new threshold intakes with respect to toxicity. Biomed Environ Sci 10:356-358.