SpringerBriefs in Physiology For further volumes: http://www.springer.com/series/10229 Klaas R. Westerterp 1 3 Energy Balance in Motion Klaas R. Westerterp Department of Human Biology Maastricht University Maastricht The Netherlands © The Author(s) 2013 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. 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The publisher makes no warranty, express or implied, with respect to the material contained herein. Printed on acid-free paper Springer is part of Springer Science+Business Media (www.springer.com) ISSN 2192-9866 ISSN 2192-9874 (electronic) ISBN 978-3-642-34626-2 ISBN 978-3-642-34627-9 (eBook) DOI 10.1007/978-3-642-34627-9 Springer Heidelberg New York Dordrecht London Library of Congress Control Number: 2012953017 v Man survives in an environment with a variable food supply. Energy balance is maintained by adapting energy intake to changes in energy expenditure and vice versa. Human energetics is introduced using an animal energetics model including growth efficiency, endurance capacity and adaptation to starvation. Animal energet- ics was the starting point for assessment of energy expenditure with respirometry and doubly labelled water and of body composition with densitometry and hydrometry. Examples of endurance performance in athletes and non-athletes illustrate limits in energy expenditure. There is a complicated interaction between physical activity and body weight. Body movement requires energy as produced by muscles. Thus, there is an interaction between physical activity, body weight, body composition and energy expenditure. Overweight is caused by energy intake exceeding energy expenditure. The questions of how energy intake and energy expenditure adapt to each other are dealt with. The evidence presented, originating from fundamental research, is translational to food production and to physical activity-induced energy expenditure in competitive sports. Another obvious and relevant clinical application deals with overweight and obesity, with the increasing risk of developing diabetes, cardiovascular disease and cancer. Finally, activity induced energy expenditure of modern man is put in perspective by compiling changes in activity energy expendi- ture, as derived from total energy expenditure and resting energy expenditure, over time. In addition, levels of activity energy expenditure in modern Western societies are compared with those from third world countries mirroring the physical activ- ity energy expenditure in Western societies in the past. Levels of physical activity expenditure of modern humans are compared with those of wild terrestrial mam- mals as well, taking into account body size and temperature effects. Taken together this book shows how energy balance has been in motion over the past four decades. Preface vii Dr. Klaas R. Westerterp is professor of Human Energetics in the Faculty of Health, Medicine and Life Sciences at Maastricht University, The Netherlands. His M.Sc in Biology at the University of Groningen resulted in a thesis titled ‘The energy budget of the nesting Starling, a field study’. He received a grant from the Netherlands Organisation for Scientific Research (FUNGO, NWO) for his doctorate research in the Faculty of Mathematics and Natural Sciences at the University of Groningen. His Ph.D. thesis was titled ‘How rats economize, energy loss in starva- tion’. Subsequently, he performed a three-year post- doc at Stirling University in Scotland supported by a grant from the Natural Environment Research Council (NERC), and a two-year postdoc at the University of Groningen and the Netherlands Institute of Ecology (NIOO, KNAW) with a grant from the Netherlands Organisation for Scientific Research (BION, NWO) in order to work on flight ener- getics in birds. In 1982, he became senior lecturer and subsequently full professor at Maastricht University in the Department of Human Biology. Here, his field of expertise is energy metabolism, physical activity, food intake and body composition and energy balance under controlled conditions and in daily life. He was editor in chief of the Proceedings of the Nutrition Society and he is currently a member of the Editorial Board of the journal Nutrition and Metabolism (London) and of the European Journal of Clinical Nutrition, and editor in chief of the European Journal of Applied Physiology. About the Author ix The content of this book is based on work performed with many students and colleagues as reflected in the references. Paul Schoffelen and Loek Wouters tech- nically supported measurements on energy expenditure with respirometry and doubly labelled water. Margriet Westerterp-Plantenga reviewed the subsequent drafts of the manuscript. Louis Foster edited the final text. Acknowledgments xi 1 Introduction, Energy Balance in Animals 1 2 Energy Balance 15 3 Limits in Energy Expenditure 37 4 Energy Expenditure, Physical Activity, Body Weight and Body Composition 47 5 Extremes in Energy Intake 63 6 Body Weight 71 7 Growth, Growth Efficiency and Ageing 83 8 Modern Man in Line with Wild Mammals 91 Appendix 97 Glossary 101 References 105 Index 111 Contents xiii ADMR Average daily metabolic rate AEE Activity-induced energy expenditure ATP Adenosine triphosphate BMI Body mass index BMR Basal metabolic rate COPD Chronic obstructive pulmonary disease DEE Diet-induced energy expenditure DEXA Dual energy X-ray absorptiometry for the measurement of body components like mineral mass EE Energy expenditure EG Energy deposited in the body during growth EI Energy intake FAO Food and agriculture organisation of the United Nations FFM Fat-free body mass FM Fat mass of the body SMR Sleeping metabolic rate TEE Total energy expenditure Tracmor Triaxial accelerometer for movement registration UNU United Nations University WHO World Health Organization Abbreviations 1 Abstract Man is an omnivore and originally met energy requirements by hunt- ing and gathering. Man evolved in an environment of feast and famine: there were periods with either a positive or negative energy balance. As an introduction to human energetics, this book on energy balance in motion starts with a chapter on animal energetics. How do animals survive and reproduce in an environment with a variable food supply? The examples on animal energetics illustrate how animals grow, reproduce and survive periods of starvation. It is an introduction to method- ology and basic concepts in energetics. Growth efficiency of a wild bird in its nat- ural environment, here the Starling, is similar to a farm animal like the Domestic Fowl. Reproductive capacity is set by foraging capacity, determined by food avail- ability and the capacity parents can produce food to the offspring. Birds feeding nestlings reach an energy ceiling where daily energy expenditure is four times resting energy expenditure. Starvation leads to a decrease in energy expenditure, where the largest saving on energy expenditure can be ascribed to a decrease in activity energy expenditure. Keywords  Activity  factor  •  Body  temperature  •  Doubly  labelled  water  method  • Energy ceiling  •  Gross energy intake  •  Growth efciency  •  Metabolizable energy  • Starvation The Energy Budget of the Nestling Starling From the late Middle Ages, nestling Starlings were harvested to prepare paté or soup. As such, Starlings were a source of animal protein in a hunter and gatherer system. Passerine birds have short incubation periods (12–14 days) and a nestling period of some weeks, characterized by rapid growth. The conversion ratio of food  to energy incorporated in the growing body is high. Here the energy budget of the nestling Starling is presented for the calculation of the growth efficiency of a wild animal in its natural environment. The result is compared with figures for the Domestic Fowl, one of our current sources for animal protein. In the Netherlands, wild Starlings were offered artificial nest sites by mount- ing ‘Starling pots’ against a building (Fig. 1.1). Pots were made from clay with a Introduction, Energy Balance in Animals Chapter 1 K. R. Westerterp, Energy Balance in Motion, SpringerBriefs in Physiology, DOI: 10.1007/978-3-642-34627-9_1, © The Author(s) 2013 2 1 Introduction, Energy Balance in Animals long neck, and a hole 5 cm in diameter as entrance. Pots were mounted against the wall of a house at a height of some meters with the neck horizontal. At the back,  against the wall, was a hole to harvest the chicks. The optimal harvest time is just before fledging, in the third week after the eggs hatch. An average brood provides four to five chicks of 70 g each or about 300 g Starling. Starlings prefer to breed in colonies. Thus, one can mount several pots on the same house. Additionally, Starlings often start a second brood, especially when taking the chicks disturbs the first brood. The Starling (Sturnus vulgaris) is a feasible subject for a field investigation. As a hole nester readily accepting nest-boxes, a Starling colony can be founded at any convenient point bounding on pastureland for foraging. The nestlings develop from hatching to fledging in 19–21 days. There is close synchrony in breeding behaviour within the colony and the adults forage in the same general area allow- ing several adults to be observed at the same time, thus duplicating observations. Growth efficiency, the relation between energy intake and the energy deposited in the body during growth, is assessed by measurement of the separate components of Fig. 1.1 Five ‘Starling pots’, mounted against the front of a house or pub, with somebody inspecting from the loft (Etching Claes Janz Visscher. The village party, 1617. With permission:  Rijksmuseum, Amsterdam) [...]... from research in animals and man is combined under the title ‘Modern man in line with wild mammals’ Chapter 2 Energy Balance Abstract  Energy balance in animals and man is a balance between energy intake and energy expenditure for body functions and physical activity Energy expenditure determines energy requirement Energy requirement is met by energy intake When energy intake does not match energy requirement,... to activities inducing an energy expenditure lower than 1.75 times BMR like sleeping, lying down, sitting and standing without movement, and 25 % of time to activities inducing an energy expenditure higher than 1.75 times BMR like standing active i.e washing dishes, walking, cycling and running (Fig. 2.9a) For total energy expenditure, the energy associated with the lower and higher intensity activities...The Energy Budget of the Nestling Starling 3 Fig. 1.2  Diagrammatic representation of the energy budget of a nestling Starling (After Westerterp 1973) the energy budget: food intake, rejecta, metabolizable energy, energy expenditure, and energy stored in growth (Fig. 1.2) Food provides the organism with energy for maintenance, temperature regulation activity and growth Of the total incoming food energy. .. fraction of energy intake Energy expenditure for food processing is 10 %, for a rat on a 1  Introduction, Energy Balance in Animals 12 diet of standard laboratory food Thus, when intake matches expenditure, energy expenditure for food processing is 10 % of total energy expenditure Energy expenditure decreases by 10 % when a rat stops eating This is the same for man, as described in Chap. 5 Activity energy. .. calculated as total energy expenditure minus the sum of energy expenditure for food processing and resting energy expenditure In the baseline situation, before food deprivation, total energy expenditure was 2 W, resting energy expenditure 1.5 W, expenditure for food processing 10 % of 2 or 0.2 W, and activity energy expenditure the remaining 0.3 W After 11 days food deprivation, total energy expenditure... Here, the main example of free ranging birds reaching foraging limits is the House Martin Energy expenditure in free ranging adult House Martins was measured while they were feeding nestlings Observations covered three subsequent years in a colony of some 20 nests at a farm where food supply was monitored continuously with a suction trap for insects at the same height as foraging House Martins of 10–15 m... capturing, injecting water and taking blood samples is minimal Some birds, as marked with regular aluminium leg rings, were observed with doubly labelled in all subsequent years Fig. 1.5  Protocol for the doubly labelled water method to measure energy expenditure in free ranging House Martins during the breeding season (With permission from Bryant and Westerterp 1980) Stable isotopes are introduced by intraperitoneal... observations in nestling feeding Swallows and Sand Martins resulted in values of 3.9 and 4.3 times resting energy expenditure, respectively (Westerterp and Bryant 1984) It confirms the energetic ceiling is reached at an activity factor of four Subsequent observations in nestling feeding Starlings resulted in activity factors ranging from 3.2 to 4.3 (Westerterp and Drent 1985) The value of the activity factor in. .. feed the nestlings Additionally the parent has to meet its own energy requirement for the activity by foraging The most energy demanding activity in this situation is flying up and down between the foraging grounds and the nest Thus, the main determinant for breeding success of the chicks is the working capacity of the parents As mentioned before, nestling feeding parents reach a ceiling that caps... gross energy, a part is voided as rejecta including both faeces and urine The remainder is commonly termed metabolizable energy Measurements of the separate components of the energy budget of the nestling Starling are described to illustrate the methodology and general principles of energetics (Westerterp 1973) Energy intake of the nestlings is measured by taking samples of the meals, and by counting . supply. Energy balance is maintained by adapting energy intake to changes in energy expenditure and vice versa. Human energetics is introduced using an. of the energy budget of a nestling Starling (After Wester- terp 1973) The Energy Budget of the Nestling Starling 4 1 Introduction, Energy Balance in Animals the main