Appetite and Food Intake Central Control, Second Edition http://taylorandfrancis.com Appetite and Food Intake Central Control, Second Edition Edited by Ruth B.S. Harris CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2017 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Printed on acid-free paper Version Date: 20161124 International Standard Book Number-13: 978-1-4987-2316-9 (Hardback) This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. 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CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging‑in‑Publication Data Names: Harris, Ruth B. S. Title: Appetite and food intake : central control / [edited by] Ruth B.S. Harris. Description: Second edition. | Boca Raton : CRC Press, 2017. | Previous edition: Appetite and food intake : behavioral and physiological considerations / edited by Ruth B.S. Harris, Richard D. Mattes (Boca Raton : CRC Press, 2008). | Includes bibliographical references. Identifiers: LCCN 2016045823 | ISBN 9781498723169 (hardback : alk. paper) Subjects: LCSH: Ingestion--Regulation. | Appetite--Physiological aspects. | Appetite disorders. | Food habits--Psychological aspects. Classification: LCC QP147 .A66 2017 | DDC 612.3/1--dc23 LC record available at https://lccn.loc.gov/2016045823 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com Contents Preface......................................................................................................................vii Contributors ..............................................................................................................xi Chapter 1 Appetite Control in C. elegans .............................................................1 Kristen Davis, Mi Cheong Cheong, Ji Su Park, and Young-Jai You Chapter 2 Central and Peripheral Regulation of Appetite and Food Intake in Drosophila .....................................................................................17 Audrey Branch and Ping Shen Chapter 3 The Hamster as a Model for Human Ingestive Behavior ...................39 Ruth B.S. Harris Chapter 4 Beyond Homeostasis: Understanding the Impact of Psychosocial Factors on Appetite Using Nonhuman Primate Models ................................................................................................67 Mark E. Wilson and Vasiliki Michopoulos Chapter 5 Untangling Appetite Circuits with Optogenetics and Chemogenetics ............................................................................91 Michael J. Krashes Chapter 6 The Use of Functional Magnetic Resonance Imaging in the Study of Appetite and Obesity ...............................................117 Selin Neseliler, Jung-Eun Han, and Alain Dagher Chapter 7 Development of Hypothalamic Circuits That Control Food Intake and Energy Balance ..............................................................135 Sebastien G. Bouret Chapter 8 Maternal and Epigenetic Factors That Influence Food Intake and Energy Balance in Offspring .....................................................155 Lin Song, Miranda D. Johnson, and Kellie L.K. Tamashiro v vi Contents Chapter 9 Monitoring and Maintenance of Brain Glucose Supply: Importance of Hindbrain Catecholamine Neurons in This Multifaceted Task ................................................................177 Sue Ritter Chapter 10 Hindbrain Astrocyte Glucodetectors and Counterregulation ..........205 Richard C. Rogers, David H. McDougal, and Gerlinda E. Hermann Chapter 11 Vagal Afferent Signaling and the Integration of Direct and Indirect Controls of Food Intake ...............................................229 Robert C. Ritter, Carlos A. Campos, Jason Nasse, and James H. Peters Chapter 12 Energy Metabolism and Appetite Control: Separate Roles for Fat-Free Mass and Fat Mass in the Control of Food Intake in Humans ........................................................................................259 Mark Hopkins and John E. Blundell Chapter 13 Pharmacotherapy for Weight Loss ...................................................277 Thomas A. Lutz and Lori Asarian Index ......................................................................................................................297 Preface Obesity is recognized as a global health issue that is associated with increased risk for chronic disease and results in increased morbidity and mortality. Excessive weight gain is a significant health issue for all westernized societies, but the United States has one of the highest rates of overweight in the world, with more than two-thirds of the adult population classified as overweight or obese. This means that the number of individuals maintaining a healthy weight is in a minority. Conceptually, it is easy to reduce body weight by lowering energy intake and increasing expenditure; in reality, weight-loss programs that depend upon dietary and behavioral changes have very low rates of success, especially over the long-term. Successful strategies can only be developed if the scientific rationale is reliable, which emphasizes the need to gain a better and more detailed understanding of the complex and integrated mechanisms that influence appetite, food choice, food consumption, and energy expenditure. During the past two decades, gastric bypass surgery has emerged as the most effec- tive treatment for obesity, but this major surgery is associated with multiple risks and emphasizes the need for less invasive treatments and strategies to reverse weight gain or to reduce incidence of weight gain in the first place. The first edition of this book reviewed knowledge on the intake of micronutri- ents and macronutrients, food choice, and opposing views on whether or not there are mechanisms that control food intake. An emphasis was placed on perspectives gained from the use of animal models versus human trials, and a large majority of the information remains current, making it unnecessary to simply update these chapters. Therefore, I have taken the opportunity with this second edition to focus on current and emerging areas of interest in the control of food intake and energy balance, spanning the advantages of new and continuously developing technologies in animal and human studies to novel perspectives on which aspect of body composi- tion is being sensed and controlled. The first section of the book is devoted to some less common models that are being used to investigate the control of food intake. Conservancy of neural circuits and neu- ropeptides across species allows the use of simple systems, such as Caenorhabditis elegans and Drosophila, to elucidate the role of specific neurotransmitters in differ- ent aspects of ingestive behavior, taking advantage of relatively simple circuitry and in-depth knowledge of the genome of the organism. At a different level, hamsters provide a rodent model for investigating neuropeptides that drive food aquisition, and the data illustrate a striking similarity between hormones and neurotransmit- ters that influence foraging and hoarding with those identified as important to con- summatory behavior, even though these two components of ingestive behavior are under independent control. In contrast to the work with the simple models such as C. elegans, group housed nonhuman primates provide a unique opportunity for test- ing the impact of social environment on food choice and consumption that is directly relevant to the human situation. Although not easily available to all investigators, it is clear that we can learn an enormous amount from animal models other than the commonly used rats and mice. vii viii Preface In the last decade, there have been major advances in technologies that facili- tate manipulation of specific central control systems. The ability to induce or knock down expression of neuropeptides in select populations of neurons in adult animals has helped eliminate the possibility of compensation for prenatal genetic modifica- tions. Acute activation or inhibition of neurons based on neuroanatomic location and neuropeptide expression with designer receptors exclusively activated by designer drug or optogentics has resulted in elegant experiments that confirm the functional and anatomic integration of different control systems. The limitations associated with the use of these invasive procedures in humans are partially compensated for by functional magnetic resonance imaging, which identifies brain areas that are acti- vated by food cues and how this is influenced by internal energy status, emotional status, and external environment. These elegant techniques have supported rapid and unprecedented progress in the identification of the neuronal circuitry that drives ingestive behavior. In parallel with the progress made at the molecular and cellular level, there has been a growing appreciation of the influence of environment on development and on long-term risk for chronic disease, including diabetes and obesity. Grounded in earlier human studies that defined the impact of severe food deprivation during ges- tation on lifelong risk for disease in the offsping, recent animal studies have explored mechanisms that mediate the effects of the developmental environment, including diet and maternal and paternal energy status, on the physiology and behavior of both the immediate offspring and of future generations. Two chapters summarize current investigations of how energy balance is influenced by epigenetics and by the impact of prenatal and neonatal environment on the development of hypothalamic circuits that influence food intake. Historically, a majority of research investigating the central control of food intake has focused on the hypothalamus, but now there is a growing appreciation for the critical role played by the hindbrain, which integrates information from higher brain areas with that coming from the periphery to determine an appropriate feed- ing response. Until recently, investigation of the long-term signals that represent the energy balance status of an organism has been independent of the study of short- term gastrointestinal and sensory signals that influence meal size and frequency. The hindbrain plays a major role in integrating these different types of information and allows changes in body energy stores to influence feeding behavior. Control of ingestive behavior is essential for homeostasis and one parameter that is critical for survival is control of blood and brain glucose. Two chapters address the association between glucostasis and ingestion. One focuses on the identification of brain areas and neuronal cell types that are glucose sensitive. The second reviews new evidence that astrocytes play a critical role in responding to changes in central glucose con- centration, representative of a new and growing appreciation for the functionality of glia in physiologic control systems. Traditionally, it has been concluded that food intake is controlled to maintain body energy stores in the form of fat. A challenge to this perspective is provided based on evidence from past and present studies that fat-free mass may drive day- to-day changes in food intake, whereas body fat correlates with intake over a period of days or weeks. It is proposed that fat-free mass is the primary determinant of Preface ix resting metabolic rate, which is the major component of energy expenditure under normal conditions and most closely predicts energy intake on a daily basis. Thus, daily energy intake is linked to fat-free mass with the opportunity for modulation by the amount of energy stored as fat. The final chapter reviews current drugs available for treating obesity, focusing on those that influence food intake and food choice with an emphasis on combination therapies. The fairly small number of approved and effective medications illustrates the difficulty in developing a pharmacological treat- ment for a highly complex system that not only integrates many different types of information associated with food-related decisions but also justifies the exploration of combining drugs with synergystic activity. It also highlights the need for further investigation of the systems that influence food intake so that less invasive, but effec- tive treatments can be developed to alleviate the reliance for body weight control on gastric bypass surgery. This intervention is not covered in this book, primarily because the restraint of food intake is due to imposition of a physical limitation on eating, rather than direct manipulation of a control system that has been implicated in maintenance of energy balance. There is, however, a large and growing literature on the physiological, psychological, and behavioral benefits and limitations of the procedure, which is providing unexpected insight into the influence of the gut on metabolism. The summary of some of the current trends in research on ingestive behavior in this edition should be considered a companion to the first edition, in which there was an emphasis on the knowledge gaps between basic and applied research. It also should provide a useful, although not exhaustive, reference for students and other professionals who need an overview of aspects of the control of ingestive behavior that are outside their area of specialization. Many of the chapters provide a historic overview of key observations that were made in a specific areas of research, which gives a valuable perspective and context not only to those chapters but also to the field in general. In addition, some of the topics addressed by the authors included in this book should increase awareness of the value of different approaches to com- mon issues encountered in the study of ingestion. As you go through the chapters presented here, it is clear that there is a significant degree of cross-reference and of reliance on similar established observations, emphasizing that no individual area within the study of appetite and ingestive behavior can stand alone.
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