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OXFORD MONOGRAPHS ON MEDICAL GENETICS

GENERAL EDITORS
Arno G. Motulsky
Peter S. Harper
Charles Scriver
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Judith G. Hall

16. C. R. Scriver and B. Child: Garrod’s inborn factors in disease

18. M. Baraitser: The genetics of neurological disorders
21. D. Warburton, J. Byrne, and N. Canki: Chromosome anomalies and prenatal
development: an atlas
22. J. J. Nora, K. Berg, and A.H. Nora: Cardiovascular disease: genetics, epidemiology, and
prevention
24. A. E. H. Emery: Duchenne muscular dystrophy, second edition
25. E. G. D. Tuddenham and D.N. Cooper: The molecular genetics of haemostasis and its
inherited disorders
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31. M. M. Cohen, Jr.: The child with multiple birth defects
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34. M. Baraitser: Genetics of neurological disorders, third edition
35. H. Ostrer: Non-mendelian genetics in humans
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44. R. A. King, J. I. Rotter, and A. G. Motulsky: The genetic basis of common diseases,
second edition
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counseling, third edition
47. I. J. Holt: Genetics of mitochondrial disease
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51. P. S. Harper: Landmarks in medical genetics
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56. P. L. Beales, I. S. Farooqi, and S. O’Rahilly: Genetics of obesity syndromes
Genetics of Obesity
Syndromes

Edited by
Philip L. Beales
Molecular Medicine Unit
UCL Institute of Child Health
Great Ormond Street Hospital for Children
London, United Kingdom

I. Sadaf Farooqi
Metabolic Research Laboratories
Institute of Metabolic Science
Addenbrooke’s Hospital
Cambridge, United Kingdom

Stephen O’Rahilly
Metabolic Research Laboratories
Institute of Metabolic Science
Addenbrooke’s Hospital
Cambridge, United Kingdom

1
2009
1
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Copyright © 2009 by Oxford University Press, Inc.

Published by Oxford University Press, Inc.
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All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording, or otherwise,
without the prior permission of Oxford University Press.

Library of Congress Cataloging-in-Publication Data

Genetics of obesity syndromes / [edited by] Philip L. Beales, Sadaf Farooqi, Stephen O’Rahilly.
p. ; cm.
ISBN 978-0-19-530016-1
1. Obesity--Genetic aspects. I. Beales, Philip L. II. Farooqi, Sadaf. III. O’Rahilly, S.
(Stephen) [DNLM: 1. Obesity--genetics. 2. Obesity--etiology. 3. Syndrome.
WD 210 G32805 2008]
RC628.G472 2008 616.3’98042--dc22
2007052866

9 8 7 6 5 4 3 2 1
Printed in the United States of America
on acid-free paper
Preface

Over the last two decades, the dissection of the genetic basis of obesity has
occupied researchers in fields stretching from anthropology to molecular
genetics. Such studies can be loosely divided into population-wide associa-
tion/linkage strategies and monogenic disease-based approaches. This book is
primarily concerned with the latter and has been conceived to showcase recent
advances in our understanding of the etiology of obesity. The contributing
authors, each a leading expert in the genetics of obesity-related syndromes,
acknowledge that in such a rapidly moving field it is not possible for a book
of this type to remain fully abreast of the research. Following large increases
in the number of obese patients now being referred to hospital clinics with
likely genetic etiologies, we perceived a need for a reference book of this type.
We have therefore set out this volume to serve as a guide to the differential
diagnosis and management of the obese patient.
The book is divided broadly into three parts: the first comprises an intro-
duction and a chapter describing approaches for assessing and investigating
the obese individual; the second describes nondysmorphic, monogenic forms
of obesity; and the third documents key multisystem obesity syndromes with
various genetic etiologies. It is as much a reference book as it is a manual and
will appeal to medical students, clinicians, nutritionists, molecular biologists,
and geneticists alike.
P.B.
I.S.F.
S.O.
v
This page intentionally left blank
Contents

Contributors, ix

Part I Introduction

1. Introduction, 3
Philip L. Beales and I. Sadaf Farooqi
2. A Practical Guide to the Clinical Assessment and Investigation
of Obesity, 25
I. Sadaf Farooqi

Part II Nonsyndromic Obesity

3. Human Leptin and Leptin Receptor Deficiency, 37

I. Sadaf Farooqi and Stephen O’Rahilly
4. Pro-opiomelanocortin Deficiency, 49
Heiko Krude and Annette Grüters
5. Prohormone Convertase 1, 63
Robert S. Jackson

vii
viii CONTENTS

6. Human Melanocortin 4 Receptor Deficiency, 81

I. Sadaf Farooqi and Stephen O’Rahilly

Part III Syndromic Obesity

7. Albright Hereditary Osteodystrophy, Pseudohypoparathyroidism,

and Other GNAS-Associated Syndromes, 91
Louise C. Wilson
8. The Clinical and Molecular Genetics of Alström Syndrome, 133
Gayle B. Collin, Jan D. Marshall, Jürgen K. Naggert, and Patsy M. Nishina
9. The Clinical, Molecular, and Functional Genetics of
Bardet-Biedl Syndrome, 147
Alison Ross, Philip L. Beales, and Josephine Hill
10. Börjeson-Forssman-Lehmann Syndrome, 187
Joanna Crawford, Michael Partington, Mark Corbett,
Karen Lower, and Jozef Gécz
11. Cohen Syndrome, 201
Kate E. Chandler and Forbes D. C. Manson
12. Prader-Willi Syndrome, 223
Rachel Wevrick
13. Syndromes with Obesity, 251
Philip L. Beales and Raoul Hennekam

Index, 279
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Contributors

Philip L. Beales Joanna Crawford

Molecular Medicine Unit Neurogenetics Laboratory
UCL Institute of Child Health and Great Department of Genetic Medicine
Ormond Street Hospital Women’s and Children’s Hospital
London, United Kingdom North Adelaide, Australia

Kate E. Chandler I. Sadaf Farooqi

Academic Unit of Medical Genetics and Metabolic Research Laboratories
Regional Genetics Service Institute of Metabolic Science
St. Mary’s Hospital Addenbrooke’s Hospital
Manchester, United Kingdom Cambridge, United Kingdom

Gayle B. Collin Jozef Gécz

The Jackson Laboratory Neurogenetics Laboratory
Bar Harbor, Maine Department of Genetic Medicine
Women’s and Children’s Hospital
Mark Corbett North Adelaide, Australia
Neurogenetics Laboratory
Department of Genetic Medicine Annette Grüters
Women’s and Children’s Hospital Institute of Experimental Pediatric
North Adelaide, Australia Endocrinology
Charité-University Medicine Berlin
Humboldt University
Berlin, Germany

ix
x CONTRIBUTORS

Raoul Hennekam Jürgen K. Naggert

Clinical and Molecular Genetics Unit The Jackson Laboratory
Institute of Child Health and Great Bar Harbor, Maine
Ormond Street Hospital
London, United Kingdom Patsy M. Nishina
The Jackson Laboratory
Josephine Hill Bar Harbor, Maine
Molecular Medicine Unit
UCL Institute of Child Health Stephen O’Rahilly
London, United Kingdom Metabolic Research Laboratories
Institute of Metabolic Science
Robert S. Jackson Addenbrooke’s Hospital
Department of Chemical Pathology Cambridge, United Kingdom
East Surrey Hospital
Redhill, United Kingdom Michael Partington
Neurogenetics Laboratory
Heiko Krude Department of Genetic Medicine
Institute of Experimental Pediatric Women’s and Children’s Hospital
Endocrinology North Adelaide, Australia
Charité-University Medicine Berlin
Humboldt University Alison Ross
Berlin, Germany Molecular Medicine Unit
UCL Institute of Child Health
Karen Lower London, United Kingdom
Neurogenetics Laboratory
Department of Genetic Medicine Rachel Wevrick
Women’s and Children’s Hospital Department of Medical Genetics
North Adelaide, Australia University of Alberta, Edmonton
Alberta, Canada
Forbes D. C. Manson
Centre for Molecular Medicine Louise C. Wilson
University of Manchester Clinical and Molecular Genetics Unit
Manchester, United Kingdom Institute of Child Health and Great
Ormond Street Hospital
Jan D. Marshall London, United Kingdom
The Jackson Laboratory
Bar Harbor, Maine
I

INTRODUCTION
This page intentionally left blank
1

Introduction

Philip L. Beales and I. Sadaf Farooqi

Never before in history has there been such an abundance of energy-rich,

highly processed foodstuffs. However, the price of progress is beginning to be
felt among more Westernized populations with seemingly inexorable rises in
the prevalence of obesity, diabetes, and cardiovascular disease. This so-called
nutrition transition combined with increasingly sedentary lifestyles is pro-
moting an obesogenic environment, which according to the World Health
Organization (WHO) is now the greatest risk factor for ill health worldwide
(Drewnowski and Popkin 1997). Latest estimates predict that 400 million of
the world’s population is now obese (World Health Organisation 2006). In
North America, the rates of overweight and obese children and adolescents
have tripled over the last 30 years. In the United States alone, one-third of the
population is obese; and this rate is climbing annually. Many countries in
Europe are closely following behind, and even less developed nations are wit-
nessing significant increases in the prevalence of obesity. This epidemic is by
no means confined to adults, with an increasing proportion of children and
adolescents becoming morbidly obese. Concomitant medical problems such
as type 2 diabetes mellitus, traditionally the domain of adults, are increasingly
recognized in children (Fagot-Campagna 2000). It is this cocktail of relative
inactivity and calorie availability that appears to be unmasking our underlying
genetic susceptibility to weight gain—presumably an advantage for many of
our “hunter–gatherer” ancestors (Neel 1962).
3
4 INTRODUCTION

Measuring Obesity

Obesity refers to an excess accumulation of adipose tissue. Exactly what con-

stitutes excessive is questionable; and furthermore, how this shall be meas-
ured is hotly debated. Adiposity is a continuous trait and is not easily measured;
therefore, a surrogate estimate of obesity now tends toward excess body
weight. This is usually presented as the weight adjusted for height or the body
mass index (BMI), calculated by weight in kilograms divided by height in
meters squared.
In 1981, Garrow first proposed a grading system based on BMI. He defined
obesity grades I, II, and III by BMI classes of 30–39.9 kg/m2 and 40 kg/m2 or
above, for both men and women. A value of 25 was considered the upper limit
of a normal or healthy BMI in North Americans and Europeans. A WHO
expert committee defined overweight as a BMI of 25 and greater and obesity
as a BMI of 30 or more (World Health Organization 1995). Extreme or morbid
obesity is described with a BMI of 40 or more.
The BMI, although easy to calculate, can be misleading if considered out of
context. For example, a short male muscle builder can have the same BMI as
an obese tall woman.

Childhood Obesity

Unlike in adults, where morbidity and mortality estimates are closely associ-
ated with BMI levels, such relationships with obesity in children are less
defined. Instead, a more statistical definition of overweight is widely used,
based on the 85th and 95th percentiles of sex-specific BMI for age in a speci-
fied reference population (Himes and Dietz 1994; Barlow and Dietz 1998). In
childhood, obesity is generally present when the BMI exceeds values in the
95th percentile for age and sex (Dietz and Robinson 2005). Although there is
no accepted definition for severe obesity in children, a BMI with standard
deviation (SD) >2.5 is often used in specialist centers. The crossing of the
major growth percentile lines upward is an early indication of risk for severe
obesity.

Obesity-Related Morbidity

Diabetes

Type 2 diabetes mellitus, traditionally the domain of adults, is increasingly

recognized in children. The first cases among children were reported in 2000
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