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Klaas R. Westerterp

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Energy Balance in Motion

Klaas R. Westerterp

Department of Human Biology

Maastricht University

Maastricht

The Netherlands

© The Author(s) 2013

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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

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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 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

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)

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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 or 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 the total number of meals per day. Meals can be sampled by the col￾lar method. Nestlings are collared with a cotton thread around the neck preventing

swallowing of a meal after feeding. Meals are removed after each parental visit

for later analysis with regard to diet composition and energy content. Depending

on age, nestlings can be collared for periods of one to three hours, between some

hours after sunrise and before sunset so as not to interfere with the very first and

last feedings of the day. The feeding frequency can be determined by automatic

counting of parental visits with an electric contact in the nest entrance. Energy

output in rejecta is measured by taking samples of rejecta, and by observing the

production frequency of rejecta. Faeces and urine are excreted together in mem￾branous sacs, an adaptation enabling the parents to remove them and thus keeping

the nest clean. The collection of samples is a simple matter, especially after the

fifth day when the nestlings automatically produce a faecal sac when handled. The

frequency of faecal sac production is determined by watching the parents as they

carry off the glistening white faecal sacs from the nest. The energy content of food

and faecal samples is determined by bomb calorimetry.

The first days after hatching, chicks are fed with spiders; subsequently: leather￾jackets (Tipula paludosa), earthworms (Lumbricidae), and beetle species comprise

Fig. 1.2 Diagrammatic representation of the energy budget of a nestling Starling (After Wester￾terp 1973)

The Energy Budget of the Nestling Starling