Table Of ContentBIOLOGY OF STRESS IN FARM ANIMALS:
AN INTEGRATIVE APPROACH
CURRENT TOPICS IN VETERINARY MEDICINE AND ANIMAL SCIENCE
Control of Reproduction in the Cow, The Elisa: Enzyme-Linked
edited by J.M. Sreenan Immunosorbent Assay in Veterinary Research and
Patterns of Growth and Development in Diagnosis, edited by R.e. Wardley and 1.R.
Cattle, edited by H. de Boer and J. Martin Crowther
Respiratory Diseases in Cattle, edited by Indicators Relevant to Farm Animal
W.B. Martin Welfare, edited by D. Smidt
Calving Problems and Early Viability of Farm Animal Housing and Welfare,
the Calf, edited by B. Hoffmann, 1.L. Mason and J. edited by S.H. Baxter, M.R. Baxter and 1.A.D.
Schmidt MacCormack
The Future of Beef Production in the Stunning of Animals for Slaughter, edited
European Community, edited by J.e. Bowman and by G. Eikelenboom
. P. Susmel Manipulation of Growth in Farm
Diseases of Cattle in the Tropics: Animals, edited by 1.F. Roche and D. O'Callaghan
Economic and Zoonotic Relevance, edited by M. Latent Herpes Virus Infections in
Ristic and I. Mcintyre Veterinary Medicine, edited by G. Wittmann, R.M.
Control of Reproductive Functions in Gaskell and H .-1. Rziha
Domestic Animals, edited by W. lochle and D.R. Grassland Beef Production, edited by W.
Lamond Holmes
The Laying Hen and its Environment, Recent Advances in Virus Diagnosis,
edited by R. Moss edited by M.S. McNulty and J.B. McFerran
Epidemiology and Control of The Male in Farm Animal Reproduction,
Nematodiasis in Cattle, edited by P. Nansen, R.l. edited by M. Courot
l<irgensen and E.J.L. Soulsby
Endocrine Causes of Seasonal and
The Problem of Dark-Cutting in Beef, Lactational Anestrus in Farm Animals, edited by F.
edited by D.E. Hood and P.V. Tarrant Ellendorff and F. Elsaesser
The Welfare of Pigs, edited by Brucella Melitensis, edited by 1.1'.1. Verger
W. Sybesma and M. Plommet
The Mucosal Immune System, edited by Diagnosis of Mycotoxicoses, edited by
F.l. Bourne 1. L. Richard and 1. R. Thurston
Laboratory Diagnosis in Neonatal Calf Embryonic Mortality in Farm Animals,
and Pig Diarrhoea, edited by P. W. de Leeuw and edited by J.M. Sreenan and M.e. Diskin
P.A.M. Guinee
Social Space for Domestic Animals, edited
Advances in the Control of Theileriosis, by R. Zayan
edited by A.D. Irvin, M.P. Cunningham and A.S.
The Present State of Leptospirosis
Young
Diagnosis and Control, edited by W.A. Ellis and
Fourth International Symposium on T.W.A. Little
Bovine Leukosis, edited by O.e. Straub
Acute virus infections of poultry, edited
Muscle Hypertrophy of Genetic Origin by J.B. McFerran and M.S. McNulty
and its Use to Improve Beef Production, edited by
Evaluation and Control of Meat Quality
1.W.B. King and F. Menissier
in Pigs, edited by P.V. Tarrant, G. Eikelenboom and
Aujeszky's Disease, edited by G. Wittman G.Monin
and S.A. Hall
Follicular Growth and Ovulation Rate in
Transport of Animals Intended for Farm Animals, edited by J.F. Roche and D.
Breeding, Production and Slaughter, edited by R. O'Callaghan
Moss
Cattle Housing Systems, Lameness and
Welfare and Husbandry of Calves, edited
Behaviour, edited by H.K. Wierenga and D.J. Peterse
by J.P. Signoret
Physiological and Pharmacological
Factors Influencing Fertility in the
Aspects of the Reticulo-Rumen, edited by L.A.A.
Postpartum Cow, edited by H. Karg and
Ooms, A.D. Degryse and A.S.J.P.A.M. van Miert
E. Schallen berger
Biology of Stress in Farm Animals: An
Beef Production from Different Dairy
Integrative Approach, edited by P.R. Wiepkema and
Breeds and Dairy Beef Crosses. edited by G.J. More
P.W.M. van Adrichem
O'Ferrall
BIOLOGY OF STRESS IN FARM
ANIMALS: AN INTEGRATIVE
APPROACH
A seminar in the CEC programme of coordination research on
animal welfare, held on April 17 -18, 1986, at the Pietersberg
Conference Centre, Oosterbeek, The Netherlands
Edited by
P.R. Wiepkema
Department oj Animal Husbandl~v-Ethology. Agricultural University
Wageningen, The Netherlands
P.W.M. van Adrichem
Department oj Animal Physiology. Agricultural University
Wageningen, The Netherlands
....
1987 MARTINUS NIJHOFF PUBLISHERS
"
a member of the KLUWER ACADEMIC PUBLISHERS GROUP
DORDRECHT / B0STON / LANCASTER
for
THE COMMISSION OF THE EUROPEAN COMMUNITIES
Distributors
for the United States and Canada: Kluwer Academic Publishers, P.O. Box 358,
Accord Station, Hingham, MA 02018-0358, USA
for the UK and Ireland: Kluwer Academic Publishers, MTP Press Limited,
Falcon House, Queen Square, Lancaster LAI lRN, UK
for all other countries: Kluwer Academic Publishers Group, Distribution Center,
P.O. Box 322, 3300 AH Dordrecht, The Netherlands
Library of Congress Cataloging in Publication Data
Biology of stress in farm animals.
(Current topics in veterinary medicine and animal
science)
Includes bibliographies and index.
1. Veterinary physiology--Congresses. 2. Stress
(Physiology)--Congresses. 3. Livestock--Behavior-
Congresses. 4. Stress (Psychology)--Congresses.
I. Wiepkema, P. R. (Pieter Reinier), 1928-
II. Adrichem, P. W. M. van. III. Commission of the
European Communities. IV. Series.
SF768.B54 1987 636.089'689 87-11326
ISBN-13:978-94-010-8(ID..2 _ e:olSBN-13: 978-94-009-3339-2
DOl: 10 .1007/978-94-009-3339-2
EUR 10604 EN
Copyright
© 1987 by ECSC, EEC, EABC, Brussels-Luxembourg.
Softcover reprint of the hardcover 1 st edition 1987
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, mechanical,
photocopying, recording, or otherwise, without the prior written permission of
the publishers,
Martinus Nijhoff Publishers, P.O. Box 163, 3300 AD Dordrecht,
The Netherlands.
v
PREFACE
This book contains the contributions to a workshop on stress
in farm animals held on April 17-18,1986 at the Pietersberg Con
ference Centre Oosterbeek. The workshop was financed by the Commis
sion of the European Communities from its budget for the coordina
tion of Community Agricultural Research and the Agricultural Uni
versity of Wageningen (The Netherlands). Its aim was to bring
together experts from different disciplines all having in common
that in one way or another they were involved in stress research.
Such a multidisciplinary encounter should not only provide an
interesting description of present day knowledge on stress, but
also promote a more integrated view on stress phenomena as they
occur in higher vertebrates. In the course of this workshop the
following fields of research were related to stress: endocrinology,
immunology, pathology, neurobiology, ethology and theoretical bio
logy. Each of these relationships was introduce& by one speaker
presenting a concise state of the art. The same relationship was
elaborated by a second speaker implementing the available knowledge
as far as possible to the farm animal situation. Therefore in this
book each discipline is represented by a duo and introduced by some
integrating remarks.
This design facilitated the exchange of relevant and often
unknown data among experts stemming from quite different fields of
research and interest. A significant fact was that many of the
attendants met each other for the first time and often had not even
heard from each other before. This book might help in bridging gaps
not only by offering the text of the different contributions, but
also by containing so many references about stress research in
different disciplines.
Of course a central question in most discussions was the
meaning of the concept of stress itself. Although it was accepted
that the concept especially refers to those forms of interaction of
the individual, in which the latter is more or less overtaxed, an
adequate answer is sti 11 ina developmental stage. Thi sis partly
caused by the existence of an impressive professional jargon. For
the main part, however, it has to do with our lack of proficiency
VI
to integrate really different view-points and data into one and the
same model. That appears to be the price specialists have to pay
for the fascinating progress they often realize in their own ter
rain.
As could be expected in our discussions, the concept had not
always the same contents. As a rule, however, it referred to the
individual response state evoked by stressors. Thanks to the
pioneering work of Mason (1971) and others we now know that such
stressors are not only physical ones - as emphasized by the founder
of stress research Selye (1936) -, but also psychological ones. In
fact the idea that the psychological aspects of stressors are
crucial is rapidly gaining ground. These neurobiological and cogni
tive aspects of stress phenomena dominated the discussions of the
present workshop. If these aspects are really crucial, they have an
enormous impact on our thinking about animal health and welfare
questions in present day husbandry systems. For stress, and es
pecially chronic stress, may significantly reduce profitable and
therefore relevant traits of farm animals, for instance, by re
ducing the efficiency of metabolism, by facilitating the perfor
mance of disturbed behaviour or by decreasing reproductive capabi
lities and the resistance against pathogens. Therefore one should
minimize stress as much as possible. To realize this we cannot
overlook cognitive and emotional processes in the organisms in
volved.
We hope that the present reflection of the workshop on stress
in farm animals may stimulate and integrate research on this most
relevant and fascinating field.
REFERENCES
Mason, J.W., 1971. A re-evaluation of the concept of "non-specifi
city" in stress theory. J. Psychiat. Res.!!., 323-333.
Selye, H., 1936. A syndrome produced by di verse nocuous agents.
Nature, 138, 32.
P.R. Wiepkema
P.W.M. van Adrichem
January 1987
VII
CONTENTS
Page
Preface V
SESSION I: STRESS AND ENDOCRINOLOGY
INTRODUCTION
ENDOCRINE ASPECTS OF STRESS: CENTRAL AND PERIPHERAL
MECHANISMS
A. Oliverio 3
ENDOCRINE ASPECTS OF STRESS: EVALUATION OF STRESS
REACTIONS IN FARM ANIMALS
J. Ladewi g 13
SESSION II: STRESS AND IMMUNOLOGY
INTRODUCTION 27
STRESS AND THE IMMUNE SYSTEM
R.E. Ballieux en C.J. Heijnen 29
EFFECTS OF BEHAVIOURAL AND PHYSICAL STRESSORS ON
IMMUNE RESPONSES
H.A. Siegel 39
SESSION III: STRESS AND PATHOLOGY
INTRODUCTION 55
PHYSIOLOGY OF STRESS: A BEHAVIOURAL VIEW
B. Bohus, J.M. Koolhaas, C. Nyakas, A.B. Steffens,
D.S. Fokkema and A.J.W. Scheurink 57
ORGAN CHANGE AND DAMAGE DURING STRESS. MORPHOLOGICAL
DIAGNOSIS
K. DHmmrich 71
SESSION IV: STRESS AND NEUROBIOLOGY
VIII
I NTRODUCTI ON 83
THE NEUROBIOLOGY OF STRESS RESPONSE CONTROL
D.M. Warburton 87
APPLICATIONS OF NEUROBIOLOGICAL STUDIES TO FARM
ANH1AL WELFARE
D.M. Broom 101
SESSION V: STRESS AND BEHAVIOUR
INTRODUCTION 111
BEHAVIOURAL ASPECTS OF STRESS
P.R. Wiepkema 113
BEHAVIOURAL RESPONSES TO STRESS IN FARM ANIMALS
F.O. Odberg 135
SESSION VI: STRESS MODELS AND THEORY
INTRODUCTION 151
THE RELEVANCE OF MODELS OF MOTIVATION AND LEARNING
TO ANIMAL WELFARE
F. Toates 153
THE PLACE OF BEHAVIOUR IN PHYSIOLOGY
M. Cabanac 187
List of participants 195
SESSION I: STRESS AND ENDOCRINOLOGY
INTRODUCTION
We have passed the stage in which the idea prevailed that the
main endocrinological response to a stressor was reflected in an
enhanced activity of the pituitary-adrenocortical system. Closely
and crucially connected with this system is the hypothalamus,
forming the so called hypothalamic-pituitary-adrenal axis. The
involvement of th~ hypothalamus also implies an important contri
bution of the autonomic nervous system comprising the sympathetic
and the parasympathetic divisions. The development of reliable
methods to measure minute quantities of hormones and of neuro
transmitters produced such a complex endocrinological picture, that
at this moment it is difficult to state which is the central or
main endocrinological response during stress.
As described by Oliverio, there is a most intricate pattern of
endocri nol ogi cal acti vi ti es evoked by ordi nary stressors. The
picture is still more complicated, as indicated by Ladewig, in that
a number of hormonal reactions change significantly over time and,
moreover, may differ from one individual to another. Apart from
this, endocrinological responses also depend on circadian rhythms
existing in all the organisms involved.
All this leads to the bewildering conclusion that neuro
endocrinological responses to stressors are so complex both in
patterns of participating hormones and in changes over time, that
final insight seems far out of reach. Although it is useful not to
overestimate scientific perspectives, we should not adhere to the
reverse. If we correct for daytime and individual factors, a most
important cue is without doubt that most if not all endocrinologi
cal events typical during stress do not occur independently. They
interact in a well-structured way. A main principle appears to be
the occurrence of feedbacks, be they negative or even positive.
Analysis of these feedback patterns may significantly improve our
understanding of how the perplexing puzzle of hormonal changes
during stress can be fitted. As indicated by Oliverio, a crucial
2
role has been reserved for the brain in controlling the endocrine
system (see also the contribution of Warburton); a specific role is
attributed to neuropeptides, among the corticotrophin relea
whi~h
sing factor (CRF) attracts special attention.
We not only have to know more about the highly complicated
interactions in the neuro-endocrine system (in which the role of
the parasympathetic system sometimes seems to be underexposed, cf.
the chapter of Bohus et a1.), but we also urgently need research
aiming at the biological significance of these endocrine inter
actions for an intact individual coping with changes in its normal
envi ronment. No doubt these endocri no1 ogi cal processes under1 i e
relevant homeostatic aspects of the organism's life. However, these
aspects not only imply classical ones like the regulation of body
temperature or of glucose availability and others, but also those
that govern homeostatic aspects of social organization and of
controllability of environmental changes. This latter point is well
illustrated in a scheme derived from Henry, indicating the re
lationship between hormonal patterns during stress and controllabi
lity of the environment. Next to these homeostatic effects, the
change in hormone profil es will al so i nfl uence the control in the
partitioning of nutrients (homeorhetic control).
The emerging picture indicates that also in farm animals
during stress very compl icated hormonal changes take place. Since
in this type of animals so-called chronic stress is common, it is
useful to remember Ladewig's remarks on chronic stress. He points
out that contrary to what the term suggests chronic stress need not
imply a continuous state of being overtaxed and upset. Although the
stressors may be present continuously, the animal's response state
needs not be a continuous one. Apart from habituation, specific
functional states (for instance, rest and sleep) may significantly
interrupt chronic stress.
