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Klaas R. Westerterp
Energy Balance in Motion
1 3
Klaas R. Westerterp
Department of Human Biology  
Maastricht University  
Maastricht  
The Netherlands
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
© The Author(s) 2013
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Preface
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.
v
About the Author
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 efi ld study’. He received a grant 
from  the  Netherlands  Organisation  for  Scienticfi  
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 Scienticfi  Research (BION, NWO) in order to work on iflght 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 efi ld 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.
vii
Acknowledgments
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.
ix
Contents
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
xi
Abbreviations
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
xiii
Chapter 1
Introduction, Energy Balance in Animals
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 efficiency • 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 
K. R. Westerterp, Energy Balance in Motion, SpringerBriefs in Physiology,   1
DOI: 10.1007/978-3-642-34627-9_1, © The Author(s) 2013