Transcript Slide 1
Tutorial: two case studies in genetics of obesity Prof. Karine Clément Inserm U872 team Nutriomique Université Paris 6/Cordelier Research Centre Endocrinology and Nutrition Dept, Pitié-Salpêtrière Hospital, Paris Two case studies in genetics of obesity Contents Slides 1. Gene discovery 3-20 2. Implication of a candidate gene in obesity 21-50 Further reading 51 Abbreviations used 52-53 BMI Curves of 3 sisters in a French family B A C 52 66 70 50 64 68 62 66 60 64 58 62 56 60 54 58 48 46 44 42 56 52 40 54 50 52 38 48 50 46 36 48 44 34 46 42 32 30 44 40 42 38 40 36 38 34 36 28 26 34 32 bmi 24 22 bmi +1 ds 30 -1 ds 28 +1 ds +2 ds - 2 ds 20 +3 ds -3 ds 18 26 -1 ds +2 ds 28 +2 ds +3 ds 22 N +1 ds 30 - 2 ds 24 bmi 32 -1 ds - 2 ds 26 +3 ds -3 ds 24 -3 ds 11416 22 S 20 16 20 18 14 18 16 16 14 12 14 12 10 12 10 8 10 8 6 8 6 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 6 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 0 1 2 3 4 5 6 7 Severe obesity form occurring very early in life Can we gather arguments for a genetic origin ? 8 9 10 11 12 13 14 15 16 17 18 Clinical observation of affected members in the French family Clinical Phenotype Biological phenotype BMI : 52, 65, 71kg/m2 Hypogonadotropic hypogonadism Early onset of weight gain Blunted growth hormone Short stature Abnormal eating behavior Central hypothyroidism Compulsive behavior Normal hypothalamic pituitary Normal RMR No pubertal development (17y) Absence of diabetes Abnormal social behavior Normal TG & CT No mental retardation secretion adrenal axis Severe obesity form occurring very early in life, with severe anomalies of food intake behavior and endocrine dysfunction Examination of the family pedigree 5 1 2 4 3 4 5 6 3 4 3 7 8 9 10 11 12 2 3 13 14 15 4 16 17 18 51 ans 27,7 19 47 ans 33 1 20 21 22 23 24 8 ans 25 12 ans 16,2 26 13 ans 68,9 27 16 ans 29,75 28 17 ans 33,69 29 19 ans 66 30 19 ans 52,61 31 23 ans 25,4 32 24 ans 27,3 Summary • Severe obesity form (early onset) • Severe anomalies of food intake • Endocrine anomalies (can also be mental retardation, sensorial deficiency…) • Consanguineous family • The origin is probably genetic • But which gene is involved? Leptin levels in the HD family 3 obese sisters 800 leptin ng/ml 3 normal weight 600 400 200 0 0 20 40 BMI kg/m2 60 80 100 Leptin levels in the HD family Sibs years height weight Leptin (ng/ml) 51y 47y 180cm 90kg 155cm 80kg BMI :27,78 BMI :33,30 145 362 12 13 16 17ans 19ans 19ans 23ans 24ans 151cm 37kg 153cm 149kg 171cm 87kg 175cm 103,2kg 154cm 165kg 159cm 133kg 173cm 76kg 160cm 70kg BMI :16,23 BMI :68,90 BMI :29,75 BMI :33,69 BMI :66 BMI :52,61 BMI :25,4 BMI :27,3 212 88 600 526 5.6 Clément, Nature, 1998 670 240 294 Hypothesis • • • • • • • Severe obesity form (early onset) Severe anomalies of food intake (compulsive behavior) Endocrine anomalies (growth and reproductive functions) (can also be mental retardation, sensorial deficiency…) Consanguineous family The origin is probably genetic Since leptin levels are higher than would have been expected based on body fat mass… …could it be a mutation in the leptin receptor gene ? LEPR is a member of the cytokine receptor family Ob-R (leptin receptor) Gp130 IL-4-R GH-R IL-12-R Mutation screening of Ob-R Strategy ? Study design T+ M P T+ Exon 14 Exon 15 T C C A T A A C A T Exon16 Ob-R SSCP polyacrylamide gel Clément, Nature, 1998 T C T A T A A C A T ......CACCAAAG gtattgtac Exon16 a Consequences of the ObRhd mutation Ž Exon 15 Exon 16 gt Exon 17 Ž Exon 15 Exon 16 at Exon 17 Exon17 Exon15 AAT AGT TTC ACT CAA GAA TGA AAA Asn Ser Phe Thr Gln Glu 826 827 828 829 830 831 Clément, Nature, 1998 * RT-PCR of Ob-R exon15⇒17 in the HD family M 300bp 200bp Clément, Nature, 1998 Leptin levels in the HD family Genotype Leptin (ng/ml) 8ans MN 51ans 47ans 180cm 90kg 155cm 80kg BMI :27,78 BMI :33,30 MN 145 MN 362 12ans 13ans 16ans 17ans 19ans 19ans 23ans 24ans 151cm 37kg 153cm 149kg 171cm 87kg 175cm 103,2kg 154cm 165kg 159cm 133kg 173cm 76kg 160cm 70kg BMI :16,23 BMI :68,90 BMI :29,75 BMI :33,69 BMI :66 BMI :52,61 BMI :25,4 BMI :27,3 MN 212 NN MM 600 MM 526 NN 5.6 Clément, Nature, 1998 MM 670 88 MN 240 MN 294 LEPR: a member of the cytokine receptor family Mutated LEPR LEPR Isoforms X X X X OBRa OBRc OBRb Cytokines 830 AAs OBRe OBRd Gp130 IL-4-R GH-R IL-12-R Following questions • Are the high leptin levels in LEPR mutation subjects related to the interruption of the leptinbrain loop? • Can we demonstrate that? Human monogenic obesities : models for biology Ob-Ra Ob-Rc Ob-Rd Mutation Human LeptinR * mutation X Ob-Rb XX Leptin receptor White adipocytes *Clément, Vaisse et al, Nature 1998 • Lack of leptin signal consequences? • Follow hypothalamic dysfunction • Measurements of leptin fractions (free and bound, leptin displacement) • Gene expression in peripheral tissue Gene expression of leptin in patients with LEPR loss of function Ob mRNA (atoml/ug) 70 60 Controls 50 Affected girls 40 30 20 10 0 35 40 45 50 55 Percent body fat 60 65 70 Clément et al, IJO 2002 Gel filtration chromatography of circulating leptin in the LEPR deficient family Wild type Homozygous Heterozygous I-125 leptindisplacement displacement I125 Leptin 1600 25 2 Ho mozyg ou s Ho mozyg ou s Hétérozyg ou s Wild typ e 20 1200 c t i o n 1400 ra 1 / F 1000 800 L ep t i n cp n m g 15 600 10 400 5 200 Fra ctio n Num ber 60 58 56 54 52 50 48 46 44 42 9 4 40 6 4 38 3 4 36 0 4 34 7 3 32 4 3 30 1 3 28 8 2 26 5 2 24 2 2 22 9 1 20 6 1 Lahlou, Clement, Diabetes, 2000 18 3 1 Fractio n Nu mber 16 0 7 4 1 1 0 0 Two case studies in genetics of obesity 2: Implication of a candidate gene in human obesity Leptin and melanocortin mutations Food intake Energy expenditure Mice Humans Ob Db 5* POMC-/- 3* Rare syndromes Adapted from D Cummings, 2003 6* fat 1* Mc4r (-/-) ? Phenotype Obese phenotype Mouse models: obesity with no other dysfunction Weight ob/ob db/db MC4R-/MC4R+/Wt Age Clement 2002 Case situation • MC4R: very good candidate gene – Melanocortin pathway (mutations in upstream genes lead to severe forms of obesity) – MC4R KO leads to severe obesity in mice – Not located in linked loci • Is MC4R implicated in obesity development ? First Questions - Which human cases (isolated vs. syndromes) - How many subjects we should screen (monogenic vs. polygenic) - Which techniques (1000 bp) Is MC4R gene an obesity gene ? • Case-control association study – Frequency in large populations and controls • Family analysis (co-segregation mutation family) • Phenotypes of subjects carrying homozygous and heterozygous mutations • Functional studies of mutated receptors Is MCR an obesity gene ? • Frequency in large populations and controls • Family analysis (co-segregation mutation family) • Phenotypes of subjects carrying homozygous and heterozygous mutation • Functional studies Association studies Adapted from Redden et al, 2007 Identifying Disease Genes Family Based Linkage Studies Members of a family Affected by the Same disease share Identical disease genes Population Based Association Studies Distribution of disease alleles is Different between Cases and Controls MC4R is an obesity gene ? (frequency) A 25 S N L H R A S Y35stop 35 D37V G G P48S 40 C Y L S 15 R N W L H L stop 10 H M T S T T11S T11A G 10 H M R G R 5 H T S H T S V N 5 N V hMC4R M hMC4R M S D V50M • In the French population : 774 subjects, 21 (2.7 %) mutations K G Y T37M G D T D I102SG T I102T 40 C S 42 Y G98R105 L 42N A C271Y C271R M200V Q S I125K FT D A Q T D D S S S D S S A S A P N V253I V C Q NP InsGT N274S PQ Y C Y C V C V F C V I I S Y 120 267 S F Y G252S C P 195 T M 281 F VT I I S I stop I Y H S 120 267 F Y IH L L C I N Q E 185 195 D 281 S M V F L N I I I N F C TI L IF F L H S L N 260 F M L E V L H V I T Q 105 F P 185 D I L NI P S D F L N Y I L F L I 201 N97D S295P S I T F N F A L I 260 P F C W L G F I V T L FMM T 127 V E V 290YL T P IS V D I 179 V SI L 201 V F F A L L V I 291 I I I 127 S 131 G T A EE 100 T I C C WV F V 290 L I C M 253 F51L S C I V 131 V ST 97 N V GF SV A MM L 179 C L T S 291 L V I A G L E N V 55 174T G C M L A A 253 100 I251L S S 97 N G LS W I S A C V F L T LG I 211M L S A I V L S NI 55 S G V C IA S A L A175T L S W 174 I S D 298 V SI L I 211 L HS V Y T II 249 I VV S137 C L P I IS C I S A168 MS L M TT L I I E L V Y P D 298 S I 168 90 D L L VL S 137 H L L I C 249 L L I G F I M M T 63 I N I S Y E A V L L 90 D P299H 216 GI A L 63 I AN V K RI A M Y I A I S L 145 S V G L F G KA I V L A V A L K RV 163 A M L R216 K242 M V S A I301T VA I 145 A M N V 163 242 A V L R I I C S LR D A N 750insGAA 312 G A 312 A VF R D V A I I C Y I Y V95I Q T L F G 70 F A H I316T R F T Q 80 Y T L F I 70 N K H T F 80 Y151 I R I N K T A 151 I T305 R M M I 305 R P M S58C 235 235 A I K I317T II L F P M K L L250W F F G I F G S N I CTCT 211 K S S N Y I S Y I K N R N R L H L S127L H A L Q A YL H H K A244E R305Q T K Q Y T 319 319 Q Q R C R C N62S K K G G E C E C E I I P E P I69M P78L A A I170V V LV L R165W 308 308 Y Y stop I137T F R165Q T150I A154Y P G231S Y R Y SR S D C SL S L L P L L G G D C L • More than 3000 subjects screened since 1998 – 2-3% of the population T5T G S D H S G Y A18C A18H N 15 L L K 35 N W E 30 S30F R 20 S S L R Y S L G 20 S S 25 S H S E 30 Y G G – – – – – Dominant transmission Early onset obesity (74% vs 57%) Variable expression in the families « common » obesity forms Heterogeneity of functional consequences Vaisse Nature genet 1998, JCI 2000 Lubrano Berthelier, 2003 MC4R mutation in human obesity Study country n Age BMI Carriers % mutants MC4R France 769 ≥18 ≥35 kg/m2 20 2.6 USA 165 18-48 ≥40 kg/m2 6 2.6 Switzerland 469 ≥18 ≥35 kg/m2 5 1.07 Spain 159 ≥18 ≥30 kg/m2 1 0.63 Denmark 750 20 ≥31 kg/m2 19 2.5 France 63 ≤18 ≥99thp* 4 6.35 UK 500 ≤18 ≥99thp 29 5.8 France 172 ≤18 ≥95thp 3 1.74 Germany 808 ≤18 ≥95thp 15 1.86 France 516 ≤18 ≥85thp 8 1.74 110 2.52% TOTAL 4371 * p = percentile Lubrano-Berthelier et al 2004 Dubern et al 2002 Lubrano-Berthelier et al 2003 Lubrano-Berthelier et al 2004 Extracellular 20 25 A N S H L R Y S S R N W 15 L H L 10 S T H M 30 5 G S C E H S H L A F* L R NH2 M T S N V S G H K G 35 Y L S S V* D Y G P G M C 40 T Y S T D D A Q S F S Q E 42 V S Y 120 P N K T T R I D S S A V I I 195 C C P 267 F Y C V Q I 185 Y I S L I F N H S M 281 H I L C I N L N L V 260 N F N D I L F L 105 T L T I V F F M Y P P S V L G I F 290 P I A I L 201 L F L I I E R W 127 S D 179 V S T T V C T M V I V F E 100 M V S* I C C C H A L T 131 N G S A A D F V P 55 G L N W 174 M L 253 V S S S S S G L I 97 V A I C I S L I C S V I L I S L S T I L S A 211 I 249 T Y L V V D 298 L I S L V I R P C M E 137 K L T N D M H Q I S 168 I 216 G 63 I T 90 L L L F G A R V V A Y I L K S V A S K 242 M I A M V 163 A I L N 145 -S L R A V A 312 V S C A T A I I D Q G F T Y R 70 K H I R F T I N L N F Y 235 Q W 80 T T I M M A 151 E I 305 R L F I P K M I F S R K C G S T K N Y N L H C I Q K E C H A T 320 R L Q Y Y E L W G I A V L P D P 308 L P S G S G L L C D Y R S S L S F L Intracellular Dubern B, Lubrano C, Clement K COOH Is MCR an obesity gene ? • Frequency in large populations and controls • Family analysis (co-segregation mutation family) • Phenotypes of subjects carrying homozygous and heterozygous mutation • Functional studies Family with severe MC4R mutation a. I 2 1 †73y >35kg/m2 78y 37kg/m2 Z=2.95 II III 2 1 58y 51kg/m2 Z=6.48 59y 28kg/m2 Z=0.73 3 35y 57kg/m2 Z=10.14 36y 23kg/m2 Z=-0.55 34y 50kg/m2 Z=8.54 IV 11y 30kg/m2 Z=6.5 100bp 200bp Vaisse C, Clément, Nature Genet 1998 24y 33kg/m2 Z=4.58 2 1 b. Clement Brest 2002 4 3 2 1 50y 28kg/m2 Z=0.87 5y 14kg/m2 Z=-1.5 Mutations in MC4R (examples) Age P, T IMC Mutation Val50Met Mutation Ser58Cys Dubern et al, J Pediatr 2001 Dubern et al, J Pediatr 2001 39 ans 100 kg, 173 cm 33 kg/m² M M Age Poids Taille Z-score de l’IMC 6 ans 38.1 kg 126 cm 5.4 DS Dubern, J Pediatr 2001 NM 35 ans 70 kg, 162 cm 26,7 kg/m² NM NM 13 ans 70 kg 165 cm 2.6 DS 11 ans 58 kg 155 cm 2.9 DS Age P, T IMC 39 ans 110 kg, 189 cm 30,8 kg/m² Age Poids Taille Z-score de l’IMC 35 ans 85 kg, 165 cm 31.2 kg/m² M NM M NM M 16 ans 75 kg 180 cm 1.1 DS 14 ans 69 kg 183 cm 0.9 DS 12.5 ans 92 kg 167 cm 4.2 DS NM 9 ans 40 kg 145 cm 1.7 DS Is MCR an obesity gene ? • Frequency in large populations and controls • Family analysis (co-segregation mutation family) • Phenotypes of subjects carrying homozygous and heterozygous mutation • Functional studies Complete loss of function leads to a severe phenotype 25 S N A L H R Y 20 S S R N W 15 L H S E 30 L S T 10 H M G R 5 H T S N V M S L G K G 35 Y S D G D T G S L 63 V I 70 A L M I 80 Y F P M S K N L L V A V C K N S V A V A I D 120 V N D I 3 y, Weight 27.4 kg BMI 27 kg/m2 Impulsivity Familial Obesity Consanguinity N I E 100 S G N 90 I I S L E N I 97 V I N L L T T L G 55 E V F V 105 T V S P F T S E 42 Q L S T Y F Q D C 40 A • • • • • L S • Homozygous deletion of MC4R gene (delAG 346-347) F H Dubern, J Pediatr, 2007 MC4R carriers and food intake behavior ? • Branson et al « Binge eating as a major phenotype of Melanocortin receptor gene mutations » NEJM 2003 • Not confirmed (Farooqi et al. 2003; Gotoda 2003; Herpertz et al. 2003; Hebebrand et al. 2004, Lubrano-Berthelier, JCEM 2006) Phenotypes of MCAR carriers Children NM M n = 59 n=4 Age (y) 11.6 ± 0.4 11.9 ± 1.5 BMI Z-score (SD) 4.6 ± 0.1 4.5 ± 0.5 Obesity early onset (y) 2.6 ± 0.3 2.5 ± 1.0 Adiposity rebound (y) 2.5 ± 0.2 3.3 ± 0.7 Fat mass (%) 43.8 ± 0.9 40.6 ± 4.7 Insulin (µUml) 20.2 ± 1.9 17.7 ± 7.2 Food Intake, JCI 2000 control n = 254 mut n = 201 mut + n=8 age (y) 52 +/- 7 42 +/- 11 39 +/-12 BMI (kg/m²) 21 +/- 2 51.5 +/- 7.5 52.5 +/- 8.0 2023 +/- 551 2393 +/- 900 2800 +/- 763 38.4 +/- 5 39 +/- 7.4 40.5 +/- 9.0 kcal/24h Lipids (%) MC4R carriers and height growth Increased Bone density Decreased bone resorption (Eleftriou Nature 2005 Ahn & Dubern Endocrinology 2006) Farooqi, NEJM, 2003 Evidence that MC4R is an “obesity gene” • Biological candidate • Invalidation in mouse models leads to obesity (KO) • Population based association studies • “Co-segregation” of genotype and phenotype in families • Loss of function of variant receptors Review in Govaerts Peptide 2005 European populations screened (>5000 obese screened, >8000 controls) UK (Farooqi &Yeo NEJM 03, Hum Mol Genet 03, JCI 00, Nature Genet 98) Finland (Valli-Jaakola, JCEM 04) France (Lubrano, Dubern, Clément Vaisse, Diabetes 04, JCEM 04, Hum Mol Genet 03, J Ped 00, JCI 00, Nature genet 98, JCEM 06) Germany (Hinney & Hebebrand, Am J Hum Genet 04, JCEM 03, Mol Psy02, Am J Hum Gent 99, Biebermann H, JCEM 06) Italy (Miraglia Del Giudice, JIO 02, Buono 05) Spain (Marti IJO 03) Switzerland (Branson, NEJM 03) Denmark (Larsen, JCEM 2005) And others… Is MCR an obesity gene ? • Frequency in large populations and controls • Family analysis (co-segregation mutation family) • Phenotypes of subjects carrying homozygous and heterozygous mutation • Functional studies Cloning of MC4R mutants 1. PCR amplification of MC4R gene from genomic DNA of patients carrying mutations; 2. Preparation of Insert and vector: 1) Insert - Digestion of purified PCR products with EcoRI / BspEI; 2) Vector - Digestion of wild-type plasmid (Pc) with EcoRI / BspEI; - Running on 1.5% agarose gel, cutting vector band and making gel extraction and purification with QIAGEN kit; 3. Ligation of vector (wt) and inserts (mutations): - Ratio of vector to insert = 1 / 5; 4. Transformation: - DH5 competent cells; 5. Miniculture; 6. MiniPrep to obtain the new plasmids with mutations (QIAGEN Kit); 7. Sequencing of the new plasmids. Assay of cAMP activity stimulated by -MSH 1. Cell culture: Human embryonic kidney (HEK 293) cell line; 2. Transfection of HEK293 cells with MC4R plasmid: - Lipofectamine with Plus reagent; - Transfection with Flag-GFP chimerical wild-type MC4R: Assay of cAMP activity stimulated by -MSH 3. cAMP activity stimulated by -MSH in MC4R-transfected cells: - Enzymeimmunoassay (EIA) with cAMP assay kit; - Transfection with wild-type or mutant MC4R (Pc): 100 cAMP (% of untreatment) 2500 80 % B/B0 60 40 2000 1500 1000 500 20 no n0 no nno 1 n10 0 Pc -0 Pc -1 Pc -1 00 M 10 M 11 M 110 0 M 20 -0 M 20 -1 M 20 -1 00 0 0 10 100 1000 10000 cAMP (fm ol/w ell) Standard curve of cAMP concentrations alpha-MSH (nM) cAMP activity stimulated by -MSH (non: non-transfected; Pc: MC4R wt; M1 & M20: mutants No. 1 & No. 20) aMSH responses Lubrano-Berthelier et al, Hum Mol Genet, 2002 125 W T 1 .0 4 ± 0 .1 6 n M V 5 0 M 1 .6 4 ± 0 .6 4 n M 100 75 S 5 8 C 1 5 .8 8 ± 7 .1 6 n M I1 0 2 S 50 25 0 -1 3 -1 2 -11 -1 0 -9 -M S H lo g M -8 -7 -6 Membrane expression Lubrano-Berthelier et al, Hum Mol Genet, 2002 N Flag MC4R GFP C Wild-type receptor V V X Ile102Ser Functional effects of obesity associated Melanocortin-4 Receptor Mutations Base Change A-307-G A-751-C 47-48insG 750 ins GATT A-31-G C-52-T C-449-T A-508-G C-493-T T-749-A T-902-C Effect on amino acid sequence Va l103 Ile Ile251 Leu 16 +12 amino-acids Frames hift at 6thTM Thr11Ser Arg18Cys Thr150Ile Ile170Va l Arg165Trp Leu250G ln Ile301 Thr Vaisse C, UCSF, 2002 NIe = normal Membrane expression Nle Nle 0 0 Nle Nle Nle Nle Nle Basal Activity Nle Nle 0 0 EC50 for MSH stimulation Nle Nle Nle Nle Nle/ C Vaisse, NAASO, 2001 Functional consequences of MC4R mutations Data from the French population AGRP N Membrane expression MSH - Intracellular retention => 56% of MC4R mutations + C X Prot Gs AC AMPc ? Food Intake Energy homeostasis Lubrano-Berthelier et al, HMG 2003 Srinivasan et Lubrano-Berthelier et al JCI 2004 Lubrano-Berthelier et al, JCEM 2006 Receptor activity Deficit of MSH response 80% of MC4R mutation Decreased basal activity => 76% of MC4R mutations Genotype-phenotype Relationships Intracellular retention associated with early onset obesity Evidence that MC4R is an “obesity gene” Further reading: three useful reviews 1: O'Rahilly S. Human obesity and insulin resistance: lessons from experiments of nature. Biochem Soc Trans. 2007 Feb;35(Pt 1):33-6. Review. PMID: 17212584 [PubMed - indexed for MEDLINE] 2: Mutch DM, Clement K. Unraveling the genetics of human obesity. PLoS Genet. 2006 Dec 29;2(12):e188. Review. PMID: 17196040 [PubMed - indexed for MEDLINE] 3: Farooqi IS. Monogenic human obesity syndromes. Prog Brain Res. 2006;153:119-25. Review. Abbreviations used I AC Acetylcholine ACTH Adrenocorticotropic hormone AGRP Agouti related peptide ARC Arculate nucleus BBS Bardet-Biedl syndrome BDNF Brain-derived neurotrophic factor CEPH Centre d’Etudes du Polymorphisme Humain CLIP corticotropin-like intermediate lobe peptide or ACTH18-39 CPE Carboxypeptidase E CPH Carboxypeptidase H CTX Collagen fragment peptide AHDGGR Dpd Deoxypyridinoline (bone resorption marker) EcoR1, BspE1 Restriction enzymes GFP Green fluorescent protein GHR Ghrelin receptor GLP Glucagon-like peptide Gs, Gi, Go Guanine binding proteins (s = stimulating, I = inhibiting) IBD Identical by descent IFT Intraflagellar transport IL Interleukin IR Insulin receptor Abbreviations used II LepR Leptin receptor LOD Logarithmic odds LPH Lipotropic pituitary hormone M Mutant MC4R Melanocortin4 receptor MSH (α-, etc) Melanocyte stimulating hormone N Normal NIe Normal NPY Neuropeptide Y PC (1, etc) Prohormone convertase 1 POMC Pro-opiomelanocortin PPARG Peroxisome proliferative activated receptor, gamma PVN Paraventricular nucleus PYY Pancreatic Peptide YY3-36 RMR Resting metabolic rate SIM1 Drosophila single-minded gene SNP Single nucleotide polymorphism SSCP Single strand conformation polymorphism TDT Transmission disequilibrium test Z-score Number of standard deviations from an age/sex adjusted mean