Applied Animal Behaviour Science, 37 ( 1993 ) 161-181 16 I

0168-1591/93/$06.00 © 1993 - Elsevier Science Publishers B.V. All rights reserved

Who needs 'behavioural needs'? Motivational

aspects of the needs of animals

P. Jensen*, F.M. Toates j

Section ~f Ethology, Department of Animal ttygiene, Swedish Universio' of Agricultural Science~
P.O.B. 345, S-532 24 Skara, Sweden
(Accepted 22 February 1993 )

Abstract

The aim of this paper is to investigate the question of whether there exist needs to perform paruc-
ular behaviours when the physiological needs of the animal are taken care of. According to current
theory, there exist apparent behavioural needs to perform parts of the behavioural repertoires of ani-
mals, fulfilling some or all of the following three criteria: ( 1 ) the behaviour patterns are mainly caused
by internal factors; (2) the tendencies are gradually built up while they are not being performed; ( 3 )
the mere performance of the behaviour patterns is rewarding. This has led to a "catalogue approach'.
where welfare theorists have attempted to divide the behavioural repertoire of different species into
those whose expression constitute needs and those that do not. We dispute this simplistic view. In
light of the data we review, we argue that it is conceptually wrong to dichotomize and rank the moti-
vational effects of external and internal factors. Motivation is inextricably a function of both, al-
though the variation in behavioural output may, in a given situation, be explained by the variation in
either. In some cases, the tendency to perform a behaviour does rise as a function of time. This may
be caused by factors associated with tissue needs or factors intrinsic to the nervous system, or both.
However, we see no reason for why a behavioural expression should be a need if its tendency builds
up with time and not so if other motivational processes are responsible for its variation. We also argue
that although species-specific motor activity sometimes might appear to be reinforcing to an animal.
this is, in most cases, difficult to ascertain. Moreover, a need may be present even if the behaviour is
not self-rewarding. In assessing the needs of animals we argue for a holistic approach to the motiva-
tional control of behaviour. Rather than focusing on one aspect, e.g. whether the behaviour is caused
by external or internal factors, we suggest that the total behavioural system is analyzed. When consid-
ering even rather simple and well-defined behaviour, like nest building of domestic sows (Sus scr~fa )
and dust-bathing of domestic hens (Gallus gallus), such a complicated picture emerges, that we re-
gard it as impossible to denote the performance of some behaviour patterns of a species as 'needs" as
opposed to other elements of the repertoire. We reject the 'catalogue approach', but not the idea that
there are needs which are best described as ethological; in the sense that preventing an animal from
carrying out a certain behaviour in a given situation might cause signs of suffering. There are probabl~
needs associated with the performance of all species-specific behaviour and those are a complex of
obtaining a goal and performing the motor patterns. Whether one wants to describe behaviour as a
need therefore depends on a knowledge of the environmental context. Thus, a behaviour may be a
called need in a particular situation.

Key words: Behavioural needs; General motivation

*Corresponding author.
~Present address: Biology D e p a r t m e n t , The Open University, Milton Keynes MK7 6AA, UK.
162 P. Jensen, F.M. Toates/Appl. Anim. Behav. Sci. 37(1993) 161-181

Introduction

When considering the welfare of animals kept by humans (e.g. on farms, in

zoos and in laboratories) the question of needs with regard to behaviour has
become a central issue. The modern discussion probably traces back to the
paper presented by Thorpe to the Brambell Committee ( 1965 ); he coined the
idea that animals possess behavioural needs apart from the more obvious
physiological needs that the behaviour serves (e.g. foraging behaviour serves
to obtain more than just physiological satiation). His suggestion was based
on the theories of Lorenz ( 1950 ), that action-specific energy would build up
inside the animal if it was deprived of the relevant releasing stimuli. So, in a
barren environment, animals would experience an internal drive for perform-
ing all those species-specific actions that were not regularly released.
When first presented, the idea of behavioural needs was presented in the
context of motivational theory. However, during the 1970s many ethologists
abandoned the study of motivation for the apparently more profitable areas
of behavioural ecology. The development of the understanding of causal
mechanisms was therefore retarded considerably. By the early 1970s so much
knowledge had accumulated within motivational theory that the over-simpli-
fied views in Lorenz's earlier writings were abandoned by most theorists (see
Hinde, 1970, for an extensive review of the state of the art at that time).
However, in rejecting the Lorenzian ideas completely, some theorists ap-
pear to have thrown out the baby with the bath-water (see for example Dawk-
ins, 1986). Many of the things that were so elegantly encompassed by the
intuitive, yet simple, 'psycho-hydraulic' model, are too often lost in modern
motivational models. For example, with regard to behavioural needs, the pre-
diction that the motor act in itself carries some motivational significance, as
in the performance of so-called vacuum activities, is often lost (Toates and
Jensen, 1991 ).
Recently, the concept of behavioural needs has experienced a revival (e.g.
Hughes, 1980; Friend, 1989; Hughes and Duncan, 1988a, 1988b). As before,
the motivational framework plays an important role. Hughes and Duncan
(1988a) explicitly state that the concept of an ethological 'need' can be clar-
ified only if it can be subsumed within the context of a satisfactory model of
motivation. Also in the German-speaking part of the ethological world behav-
ioural needs have been important issues of debate (Tschanz, 1987 ).
We have argued before that features of the Lorenzian theory - - homeostatic
models, the state-space model and models of goal direction - - should all be
included in a comprehensive discussion of animal motivation aiming at un-
derstanding the needs of animals in captivity (Toates and Jensen, 1991 ). We
identify a need as a state, which if not attained causes suffering to an animal
as indexed by disturbed behaviour, an increased risk of pathology a n d / o r a
hormonal profile consistent with stress (e.g. abnormal levels of hormone se-
P. Jensen, F.M. Toates/Appl. Anita. Behav. Sci. 37 (1993) 161-181 l 63

cretion). Such states clearly exist in the case of lack of nutrition or abnormal
body temperature. Can the definition be extended to include states that exist
when an animal is unable to perform a particular behaviour, but physiological
needs are taken care of?. This is the question we set out to investigate in this
paper.
We will extend our view on current motivational theory, as expressed in the
approach of Toates and Jensen ( 1991 ), to its relation to the issue of the needs
of animals in captivity. In doing so we will, to some extent, concentrate on
behavioural systems that are in the focus of the welfare debate and which we
believe at the same time illustrate our thesis.

The behavioural system

In Fig. 1 the model diplayed simply brings together those possible pathways
through which external and internal causal factors may interact with the be-
haviour of the animal. Both internal and external causal factors affect the
motivational state of the animal. Of course, the same causal factors may af-
fect different motivational systems in different ways (e.g. the tendency of a
smooth newt to breathe is affected not only by the depletion of the oxygen
reserves, but also by causal factors relevant to sexual behaviour; Halliday,
1977 ). Furthermore, external causal factors may affect the level of those that
are internal. For instance, exposure to certain stimuli may alter the level of
various hormones; as when oestrus is synchronized in a group of female pigs
(Hemsworth, 1982). Also well-known is the way nesting in canaries is orga-
nized; external stimuli cause hormonal changes which subsequently alter the
responsiveness of the birds to other stimuli (Hinde and Steel, 1972 ).

--~" factors
External 11 I
Internal
Comparison ~ factors

Goal
(set-point)

v Motorprogram
Fig 1. A general model of a behavioural system, integrating the possible pathways by which
behaviour could be controlled. The model is further explained in the text.
164 t'. Jensen, F.M. Toates/AppL Anim. Behav. Sci. 37 (1993) 161-181

The motivational state is a state of the nervous system. It determines the

likelihood that an animal will engage in a certain behaviour, or the strength
of its tendency in competing for expression. This state may include a goal
representation. When it exceeds a certain value, the relevant behaviour pro-
gram will be released and the consequences of it will feed back on the goal
representation (i.e. the continuous discrepancy between the state of the en-
vironment and the set point is evaluated), on the external causal factors (e.g.
a rival is removed from a territory), on the internal causal factors (e.g. through
a build-up of energy stores during feeding) and directly on the motivational
level (e.g. through proprioceptive feedback). Naturally, the prevailing exter-
nal factors may control and guide the motor program, causing flexibility in
behavioural output. Of course, all the relationships in the model are affected
by individual, genetic and experiential factors. The different pathways of
feedback outlined in the model are, therefore, the possible ways in which a
specific motivation could be turned off, or at least lowered to a level where it
can no longer compete with other behavioural tendencies.
As we will attempt to show in this paper the discussion ofbehavioural needs
is often limited to some specific aspect of the control of the behaviour, e.g.
whether it is caused mainly by internal causal factors. In our opinion scien-
tific understanding of the needs of an animal, for example under barren con-
ditions in captivity, must be based upon detailed examination of all the path-
ways outlined in the model for the different behaviour systems that are of
interest. For a specific behavioural system we need to know how internal and
external factors interact under different conditions; how the motor program
is designed with regard to sequencing and flexibility; how the feedback pat-
terns work. As we will show further on, it is only in this way that a compre-
hensive understanding of behavioural needs is possible.
Since the theory ofbehavioural needs relies heavily on motivational theory,
we will continue with a review of some frequently cited models of motivation.
This will serve to substantiate our arguments that a holistic approach is nec-
essary and that this can be obtained only by integrating features of different
motivational approaches.

Motivational models

In the influential model of Konrad Lorenz (1950) action-specific energy

accumulates as a function of time, represented by flow of water into a tank.
The height of the fluid is analogous to the strength of drive and, as fluid ac-
cumulates, the pressure on a valve increases. A releasing stimulus, repre-
sented by a weight, exerts a tendency to open the valve from outside, when
the valve opens and fluid flows out this symbolizes the overt behaviour. A
feature that has tended to be forgotten in later presentations of the model is
P. Jensen, F.M. 7bates/Appk Anim. Behav. Sci. 37(1993) 161-181 165

the original explicit incorporation of positive feedback, leading to a certain

persistence of a released behaviour.
Later, Lorenz modified his original model, removing the dichotomy be-
tween the action of internal and external factors (Lorenz, 1978). Instead of
representing external causal factors with weights, they were now symbolized
by fluid poured into the tank. In this v~ :sion of the model all pressure on the
valve works through the fluid level.
If we leave aside the energy-based mode of action of the models some gen-
eral suggestions about the causation ofbehaviour can be deduced from it:
1. A behaviour is released by a combination of internal and external factors.
2. When a behaviour is performed, after an initial period of positive feed-
back, the motivation will gradually wane and the behaviour stop, regardless
of the consequences of the behaviour.
3. On non-performance of a specific act the motivation for that act gradually
increases.
Basically, this is a variety of a homeostatic model in that it keeps an inter-
nal parameter within certain limits. The principle of homeostasis is widely
used in psychological models, e.g. to explain the regulatory basis of feeding
and drinking (McFarland, 1974).
Some authors have dichotomized motivations into those fitting homeosta-
sis, e.g. feeding, and those not doing so, e.g. sex (Grossman, 1967). It was
argued that the former class was internally driven while the latter was exter-
nally driven. Others have argued that incentive motivation theory could ex-
plain a wider number of results and would enable sex and aggression to be
discussed in the same context as feeding and drinking (e.g. Toates, 1986 ).
In the applied branches of ethology dealing with animal welfare, the model
of Wiepkema ( 1987 ), has become influential. According to this, animals con-
tinuously monitor the state of the outer world (Istwert) and compare this to
a Sollwert (an internal representation of the desired state). When there is
discrepancy action is taken to decrease it. As long as the action causes the
discrepancy to decrease the animal experiences a feeling of pleasure. This
model bears some similarities with both homeostatic and incentive-based
models and closely resembles the model of Deutsch ( 1960 ).
As with the Lorenzian model, some general things can be deduced from the
homeostatic models (including the Wiepkema model ).
1. Behaviour is released when a value of a controlling variable departs from a
set-point, an equilibrium optimum.
2. The behaviour turns motivation off by reducing the discrepancy of the
compared values.
3. When there is no departure of the measured variables from their set-point
values, the behaviour is not released.
We see that the models have some similarities. Both the Lorenzian and the
homeostatic approaches claim that behaviour is released as a consequence of
166 P. Jensen, F.M. Toates/AppL Anita. Behav. Sci. 37 (1993) 161-181

the departure of some variable from a certain value and turned off when the
value is set back again. However, the differences are significant. While Lorenz
claimed that the actual performance of the motor acts is the essential feed-
back mechanism, the homeostatic models generally claim that it is the con-
sequences of the behaviour that are relevant for turning off motivation (this
is also true for the Wiepkema model). Furthermore, Lorenz stated that the
motivational state will not remain constant in a constant environment. Mo-
tivational levels will gradually rise as a consequence of the non-performance
of specific acts. An animal following, for example, the Wiepkema model would
not necessarily experience a change in internal factors as a function of time
since last performance (although the 'Sollwert' might very well fluctuate with
levels of internal factors).
Hughes and Duncan (1988a) presented a model specifically aimed at in-
corporating the needs-construct within a motivational framework. According
to their view motivation is affected by internal factors while the behaviour
output is modulated by the perception of external stimuli. Both appetitive
and consummatory behaviour exert a positive feedback on motivation to start
with and only later is the feedback negative. Also the achievement of func-
tional consequences acts as a negative feedback on the motivation level and,
via the change in external stimuli, on the behavioural output.
The model resembles both the Lorenz and the Wiepkema approach. Like
Lorenz, Hughes and Duncan put much emphasis on the proprioceptive feed-
back from actually performing the motor patterns, but they also, like Wiep-
kema, recognize the effect of the consequences of the behaviour. As in the
early version of the Lorenzian 'hydraulic' model, motivation is seen as a dis-
tinctly internally caused state, something that Lorenz changed in his later ver-
sion. Although it would be possible to include appetitive behaviour in the
model of Wiepkema (Wiepkema, 1987), Hughes and Duncan are more ex-
plicit in stating that a phase of appetitive actions will precede a coi~summa-
tory act.
In conclusion, we see that the different models tend to emphasize slightly
different aspects of the control of behaviour that is outlined in Fig. 1. We will
argue for a more holistic approach to the understanding of the needs of ani-
mals which considers all the pathways of Fig. 1. However, before going into
detail, we need to examine earlier ways of dealing with the issue.

Approaches to 'behavioural needs'

What is a 'behavioural need'? Even though the concept is widely used dif-
ferent researchers appear to have different opinions about its precise mean-
ing. In a workshop held in London 1983, but not published until 1988, there
were great difficulties in reaching a consensus on simple statements about the
concept. Apart from a general formulation on different kinds of needs that
P. Jensen, F.M. ToatesL4ppL Anim. Behav. Sci. 37(1993) 161-181 167

behaviour may serve and the relation between normal behaviour and health,
the closest the workshop came to a definition was the statement: "Animals
have behavioural needs, which arise when an animal is motivated" (Hughes,
1988 ). Since an animal will probably never experience the state of being non-
motivated this definition does not clarify much, but rather raises some com-
plex issues.
It appears to us that 'behavioural needs' is generally used to describe the
need to perform a specific behaviour pattern whatever the environment is
like and even if the physiological needs which the behaviour serves are ful-
filled. It is also implicit that the performance ofbehaviour constituting needs
are motivationally different from other behaviour in the repertoire of an an-
imal. Since animal welfare, according to this view, is threatened when the
needs cannot be met, it becomes essential to distinguish the behavioural needs
from other behaviour (or the 'necessities' from the 'luxuries'; Dawkins, 1983,
1990; Matthews, 1991 ). This leads to a 'catalogue approach', where the aim
of the research is to produce a species-specific list of behavioural expressions
that have to be allowed for when an animal is kept in captivity. We will at-
tempt to show that such an ambition is fruitless and will help the welfare issue
very little.
Controversies surrounding the concept of 'behavioural needs' are closely
linked to what motivational model one prefers. On the one hand, starting
with Thorpe (Brambell, 1965 ), there are those who regard the Lorenzian ap-
proach as being the most fruitful way to look at motivation. They explicitly
rely upon the idea that an internal build-up of drive for specific acts and be-
lieve that the increased tendency to perform certain behaviours is experi-
enced by animals as latent needs to perform them (e.g. Friend, 1989 ). On the
other hand we find authors rejecting the existence of an internal build-up of
motivation, relying instead on models like those of Wiepkema. Along this line
an influential paper was published by Baxter ( 1983 ) who coined the concept
of'psychological needs' as an alternative, meaning that the animal has a goal
for its activities and if the goal can be obtained without the animal having to
take action the behaviour will never be released.
These may be regarded as two extreme positions along a line, where the one
side focuses on the internal causal factors and the other on the external. In
between we find attempts to search for a more balanced approach. Hughes
( 1980, 1988 ) suggested a model for behavioural needs based on the degree of
contribution of internal factors to the motivation of certain behaviour. Ac-
cording to this there are certain activities which are almost entirely dependent
upon fluctuations in internal factors for their performance, e.g. feeding, nest-
ing and pecking in hens, while others depend more exclusively on variations
in external factors, e.g. agonistic behaviour and anti-predator responses. Along
a gradient other behaviour patterns are controlled by various combinations
of external and internal factors, e.g. dust-bathing in hens. Hughes is of the
168 p. Jensen, F.M. ToatesL4ppl. Anita. Behav. Sci. 37 (1993) 161-181

opinion that behavioural needs are found among those behaviour patterns
which are controlled mainly by internal factors.
This opinion is more explicitly held by Hughes and Duncan (1988a):
"When behaviour is normally elicited by the presence of external factors then
the question of ethological 'need' obviously does not arise; if such a phenom-
enon as 'need' does exist then it will appear in the form of behaviour patterns
motivated mainly by internal factors".
In general, the various approaches described above portray the present state-
of-the-art opinions on the concept. Accordingly, a behavioural need would be
to perform a behaviour pattern fitting some or all of the following criteria:
1. It is mainly caused by internal causal factors.
2. Its tendency is gradually built up while it is not being performed.
3. The mere performance of the behaviour pattern is rewarding; the animal
will attempt to perform the motor patterns regardless of the state of the
environment and of the functional consequences of the activity.
A behaviour fulfilling these criteria would be expressed regularly, irrespec-
tive of whether it is needed for obtaining any functional goal, e.g. making an
animal satiated or achieving a nest. The need to perform the behaviour would
also be independent of the type of environment in which the animal lives.
Prohibiting the performance of them would be associated with negative wel-
fare. However, prohibiting other behaviour patterns, not fitting the criteria,
might be safe from a welfare point of view.
In the remainder of this paper, we will dispute this (in our view) simplistic
approach. When we start to look in detail at even some seemingly simple be-
havioural systems we find that such a complicated picture emerges that it
must be misleading to categorize them into classes dependent on, for exam-
ple, an assumed dichotomy between external and internal factors. As will be
seen this leads us to the conclusion that it is not possible to classify some of
the behavioural expressions of animals as needs in contrast to others. We will
examine the three criteria above in some detail, in order to show that each of
them are too simplified to be useable in a general theory on behavioural needs.

Can external and internal factors be dichotomized?

Is motivation more or less internally caused? This is a discussion which can

be as misleading as arguing whether a behaviour depends more upon genetics
or the environment (Hogan and Roper, 1978 ). We have suggested previously
that it is conceptually wrong to attach weightings of importance to such fac-
tors (Toates and Jensen, 1991 ). However, it is possible to discuss the source
of variation in behaviour in a given situation. For instance, feeding as well as
aggression depend upon both external and internal factors; in a certain situa-
tion the variance in each may be explicable in terms of variations in either
factor. Feeding is often assumed to be a typical behaviour controlled by inter-
P. Jensen, F.M Toates/AppL Anim. Behav. Sci. 37 (1993) 161-181 169

nal factors. Consider, for example, a rat in a standard and undisturbed labo-
ratory cage with food ad lib. The variations in feeding behaviour in this ani-
mal might indeed be explained almost entirely in terms of variations in its
internal state. However, at other times feeding might be initiated in a 'sa-
tiated' rat by presenting a cue previously associated with food presentation
(Weingarten, 1984 ). A 'satiated' pig may start feeding when seeing other pigs
eat (Hsia and Wood-Gush, 1983 ), through the process of social facilitation.
In these cases, variability in behaviour could be attributed largely to changes
in the external world. In the classic stress studies of Mason ( 1975 ) on mon-
keys it was found that stress, as indexed by an elevation of corticosteroids,
was not a consequence of food-deprivation per se, but the combination of
food deprivation and seeing other monkeys eating. Fruit-flavoured, non-nu-
tritive placebo food prevented the rise in corticosteroids that would normally
be associated with food deprivation.
Aggression is often believed to be a typical representative ofbehaviour con-
trolled by external factors. However, the aggressive behaviour of territorial
birds may fluctuate in short cycles, the birds switching between singing and
fiercely driving away intruders, followed a few minutes later by feeding close
to other birds of the same species (Hinde, 1970). In this case, the behaviour
obviously depends on variations in the internal state of the animal.
The scale developed by Hughes (1980, 1988), running from external to
internal weighting of responsibility, might therefore be misleading. One might
conceive it in terms of the normal source of variability in a given situation,
but it then becomes situation-specific and not able to generalize. It would not
enable one to list the behavioural needs of a given species that would be valid
in all environments.
It is also necessary to consider the source of variation in the internal con-
tribution to motivation, which may be very different. This was also recog-
nized by Hughes (1980). Internal factors may vary because of tissue needs
(e.g. temperature control), or from factors intrinsic to the nervous system
(e.g. in dust-bathing), or a combination of both of these (for example feed-
ing). The type of internal factors that are working in generating a certain mo-
tivational state may be of great importance in welfare issues.

Example 1: Nest building in sows

An interesting example is provided by nest building in sows. Like their

ancestor, the wild boar, sows in free-range conditions will construct a nest
during the last day of gestation in which they will give birth to their young
(Gundlach, 1968; Martys, 1982; Jensen, 1989 ). Since sows in intensive hous-
ing systems are often confined or tethered in very barren pens during farrow-
ing, there is much concern about whether sows actually need to perform nest-
ing during this period (e.g. Baxter, 1983 ). According to the theory of Hughes
170 t", Jensen, F.M. Toates/AppL Anim. Behav. Sci. 37 (1993.1161-181

( 1988 ) this could be solved by examining whether the behaviour is mainly

attributable to internal or external factors. But the picture is far more compli-
cated than this.
Nesting in sows starts approximately 10-20 h before the onset of farrowing
(Jensen, 1993 ). This coincides with a sudden rise in plasma prolactin levels
(Vale and Wagner, 1981 ). Nesting can also be induced by an injection of
prostaglandins (PGF-2-alpha) and in a series of experiments Widowski et al.
(1990) showed that the only hormonal change that was correlated with both
spontaneous and induced nesting was the prolactin rise. It can therefore be
concluded that the rise in prolactin level may be one important causal factor
involved in inducing nesting.
In an attempt to distinguish the effects of internal and external factors in
controlling the sequencing of behaviour, Jensen ( 1993 ) studied the perform-
ance of nesting activities in sows in pens that were kept free from relevant
stimuli (absolute bare concrete pens) compared with sows in similar, but en-
riched pens which contained a soil bed to root in and straw in a basket to
collect nest material from. No differences in the amount of nesting activities
were found between the groups. However, the sows in the impoverished pens
performed only the behaviour patterns that belong mainly to the first part of
the nesting sequence (nosing, rooting and pawing) and did not attempt to
carry out any vacuum gathering or arranging.
The results clearly indicate that there is no unitary 'nesting drive', since
different sequences of the behaviour are differently affected by experimental
manipulation. 'Nesting', without further specification, cannot therefore be a
need according to the criteria outlined previously. Of course, this may mean
that nesting is a complex of two different motivational processes: one being
mainly motivated by internal factors and one depending on external. Perhaps
then the first part of the nesting (nosing, rooting and pawing) would qualify
as a need sensu Hughes. However, there is evidence that external factors may
be as important in inducing the behaviour under particular circumstances.
Jensen (1988) described some cases of sows in a semi-natural environment
that were severely disturbed after having finished the nest (e.g. by ants ), and
that left the nest and recommenced the nest-building process on a new site.
Changes in hormonal levels were of course not likely to have caused this sec-
ond onset of nesting.
A more likely interpretation is that the increase of prolactin triggers a mo-
tivational state where nest-building tendency dominates. This state is then
associated with a goal representation and at the same time triggers the first
part of the motor program. If the sow does not achieve something which is
satisfactory, which decreases the difference between Istwert and Sollwert, in
the terminology of Wiepkema ( 1987 ), the next part of the motor program is
not started. If the sow has proceeded through all the motor program and
achieved a nest, but there is still a mismatch, for example because the nest is
P. Jensen, F.M. Toates/Appl. A nim. Beha v. Sci. 37 (1993) 161 - 181 I7 1

soaked, or full of ants, the remaining discrepancy may cause the behaviour to
return to the start-point. The question of what factors are responsible for elic-
iting the behaviour is thus one of determining the amount of variation in
behavioural output that is attributable to the variation in different factors
under specific environmental conditions. Nesting in sows clearly reflects a need,
but it would be wrong to say that the need is only to perform a certain set of
motor patterns in any environment.
The fact that sows when prevented from nesting before farrowing develop
abnormal behaviour, such as stereotypies, (Baxter, 1982) does indeed indi-
cate that the behaviour is associated with a need. Further, sows kept confined
at farrowing and thus inhibited from nesting have higher levels of various
pathologies at delivery (B~ickstr~Sm, 1973 ), which is also in accordance with
the idea of a need in connection with nesting (however, general confinement
effects may of course also be of importance). But we regard it as fruitless to
attempt to determine whether nesting as such is more caused by internal than
external factors.
The same difficulty in dichotomizing motivation in relation to nesting is
well-known from other species. For example, Hinde and Steel (1972) found
that in canaries the performance of different parts of the nest-building se-
quence is affected by the stimuli encountered during earlier phases. It is there-
fore difficult to regard the behaviour output as either internally or externally
motivated. Even a comparatively simple vertebrate, like the cichlid-fish
Lamprologus ocellatus, shows a remarkable flexibility with regard to building
activities. This species uses old, empty snail shells as breeding nests, digs the
shells into the sand and then covers them with substrate. The length of the
different phases has been found to depend on the stimulus situation; for ex-
ample an artificial sand pit shortened the digging, or caused it to disappear
completely, while enlarging the inside of the pit lengthened the oversanding
phase (Haussknecht and Kuenzer, 1990). These findings support the idea of
a goal representation controlling at least some aspects of the behaviour.

Example 2: Dust-bathing in fowl

Similar problems occur when we consider the dust-bathing behaviour of

poultry. This is a characteristic behaviour pattern of the species and since
battery hens are prevented from performing it, it has become a central issue
in the welfare debate on poultry (e.g. Duncan, 1978; Dawkins, 1983). The
behaviour is performed with a diurnal rhythm, showing peaks in the after-
noon. During the behaviour the hens go through a sequence of motions caus-
ing sand, earth or dust to 'shower' through the plumage and thereby reduce
the lipid contents of the feathers (Borchelt, 1975; Vestergaard, 1982 ). Under
normal conditions, the dust-bathing bouts vary between 5 and 30 min, and
there may be large intra-individual variations from one day to the next. The
172 P. Jensen, F.M. Toates/Appl. Anim. Behav. Sci. 37 (1993) 161-181

pronounced diurnal pattern of the behaviour has led some researchers to as-
sume that diurnally fluctuating internal factors are the important sources of
motivation (Vestergaard, 1982). This gains some support from an experi-
ment in which uropygial-gland-extirpated birds performed no less dust-ba-
thing than intact hens, in fact they performed slightly more (N~nrgaard-Niel-
sen and Vestergaard, 1981 ). Since the main source of feather lipids is the
uropygial gland, the results indicate that lipid accumulation (i.e. an external
factor) is not a major causal factor inducing dust-bathing.
However, this is only part of the truth. In detailed studies, Van Liere et al.
( 1991 ) found that during normal oiling behaviour - - the act whereby the
lipids from the uropygial gland are distributed into the plumage - - hens place
most of the oil on the breast feathers. In another experiment, the same au-
thors applied fresh and stale lipids mainly to the breast feathers during a sand
deprivation period and studied the effect on subsequent dust-bathing. The
administration of stale lipids caused an increase of 12% in the duration of the
first dust-bath following deprivation, compared with a control treatment
without lipids (fresh lipids hardly affected the behaviour at all). Within the
dust-bath, the effects were even more pronounced: the total number of side-
lying and side-rubbing elements doubled and tripled, respectively. These re-
sults show not only that the initiation of dust-bathing seems to depend on
external factors as well as internal, but also that external factors may affect
different elements of the dust-bath sequence in different ways. Again, this
indicates that it is not possible to break down the question of needs associated
with performing dust-bathing to a simple question about whether the behav-
iour is internally or externally motivated - - it is both, and the contribution of
each type of factor will vary with prevailing environmental conditions.

Does tendency build up with time?

There is a clear difference between, on the one hand, postulating that the
variation in performance of a certain behaviour is dependent mainly on vari-
ations in internal factors and, on the other hand, assuming that the internal
factors increase in strength in the way assumed in the Lorenzian model. This
difference is perhaps not always realized. For example, Hughes and Duncan
(1988a) state: "For any behaviour pattern which is largely governed by inter-
nal factors, motivation levels will sooner or later increase above threshold"
(see also Duncan and Poole, 1990). Other authors again simply take it for
granted that such time-dependent processes occur (e.g. Friend 1989; Dell-
meier, 1989 ). However, there is no reason why this has to be so and any state-
ment on the issue should rely on specific empirical data. The strength of in-
ternal factors does not have to increase as time passes; it could even decrease,
or be continuously suppressed by the action of other causal factors on com-
peting motivational systems (e.g. Hinde, 1970). In those cases, the expres-
P. Jensen, F.M. Toates/AppL Anim. Behav. Sci. 37(1993) 161-181 173

sion of behaviour being controlled in this way, would not necessarily consti-
tute a need according to the requirements outlined earlier. Whether motivation
increases with time is therefore a different question than whether the varia-
tion in behavioural output is explicable by the variation in internal factors.
In some cases there is clear evidence for time-dependent increments in mo-
tivation. Indeed, the tendencies to drink or feed increase as a function of the
time since the behaviour was last performed. However, this can mainly be
ascribed to metabolic factors and water loss; phenomena that are better ex-
plained by classical regulatory physiology than by the Lorenzian postulations
of action-specific energy. Probably extra-neural physiological changes can ex-
plain much of the increase in behavioural tendencies with time. In that case,
a need to perform some specific motor acts might never arise.
In other cases, where the behaviour is not normally associated with a cor-
responding extra-neural variable, there appears to be a considerable variation
in the tendency for behaviour to appear at regular intervals. In regard to
aggression, although it depends to a large extent on variations in internal fac-
tors (like testosterone levels), there is little evidence that the motivation in-
creases as a function of time since last performed (Archer, 1988). However,
other systems do seem to programme behaviour at intervals in an unchanging
environment. Rodents make regular patrols of their environment (Ely and
Henry, 1978). The activity of small nocturnal rodents in a running wheel
seems to be intrinsically programmed (Kavanau and Richter, 1968 ). Preda-
tion is motivationally distinct from aggression and feeding. In some cases it
occurs out of its normal context and appears to increase in tendency as a func-
tion of time since last performed (Polsky, 1975). Dust-bathing in hens and
quail also increases in tendency as a function of time (Vestergaard, 1980;
Schein and Statkiewicz, 1983) and Nicol (1987) found a rebound in the fre-
quency of performing such behaviour as wing-stretching and wing-flapping
after a period where the hens could not carry out these behaviour patterns.
Therefore, in some cases there may clearly be an increase with time in ten-
dency to perform certain behaviours not caused by extra-neural factors only.
This may even lead to the performance of vacuum activities. For instance,
hens in battery cages frequently perform dust-bathing movements on the wire
floor (Black and Hughes, 1974). However, it is not clear what neural factors
mediate this tendency. It might be a 'drive', that will lead the animal to per-
form the activities regardless of the state of the outer world, or it may be an
increased sensitivity to stimuli. The latter opinion is largely favoured by Hinde
(1970). On the other hand, some of the data referred to by Hinde lends sup-
port to the view that deprivation may specifically increase the urge to per-
form, for example, food-searching behaviour.
In the classical studies of nest building in canaries (Hinde and Stevenson,
quoted by Hinde, 1970), some results may indicate that time-dependent ac-
cumulation of tendency is part of the explanation of how the behaviour is
174 P. Jensen, F.M. Toates/AppL Anita. Behav. Sci. 37 (1993) 161-181

sequenced. Periods of gathering nest material are followed by building pe-

riods. The length of each period depends on the length of the previous in a
way compatible with the idea that the tendency to build gradually increases
as the bird gathers material, finally out-competing gathering and vice versa.
So, even if there are cases where behaviour tendency obviously increases as a
function of time, if one wishes to understand the needs of animals, it is nec-
essary to distinguish the causes of such an increase.
There is, however, no reason why the need to perform a specific behaviour
should be stronger if it is triggered by a time-dependent motivational process
than if it is released by factors prevailing in the environment. For example, a
battery hen may not experience any need to carry out anti-predator avoidance
just because a certain time has elapsed since it was last performed. But a per-
son (even the caretaker) entering the stable may be conceived as a potential
danger and cause a sudden increase in flight tendency. The need to perform
the behaviour once released therefore does not depend on the immediate mo-
tivating factors.

Is the performance of behaviour rewarding?

According to the models of Lorenz, and of Hughes and Duncan, the per-
formance of motivated behaviour is in itself an important factor in eventually
decreasing motivation and turning behaviour off. This is very similar to the
statement that the behaviour in itself is reinforcing, has affective value or is
hedonically loaded (Epstein, 1982 ). For example, Hughes et al. (1989) sug-
gest that the performance of nest-building behaviour, regardless of its conse-
quences, is reinforcing and is thus important to the fowl. However, the actual
meaning of such suggestions is often not clear.
While it is commonly agreed that such external factors as food, water and
sexual contact can have hedonic, incentive or affective value (e.g. Cabanac,
1971 ), motor performance has usually been seen as hedonically neutral,
meaning that the animal is moved into action by the pull of incentives or
associations with them (Herrnstein, 1977). However, other evidence may
also be interpreted to show that certain response patterns are particularly as-
sociated with certain motivational states (Gallistel, 1980) and behaviour
performance in itself plays a role in the reinforcement process (Glickman and
Schiff, 1967; Herrnstein, 1977 ).
Traditionally in psychology reinforcement has been used to describe a pro-
cess of strengthening a response that underlies learning. Exactly what reinfor-
cer can reinforce what response (Bolles, 1970) and how the mechanism of
reinforcement actually works (Bindra, 1976) have been topics of lively de-
bate, but the concept of strengthening a tendency is implicit throughout.
Sometimes, the observation that animals will also tend to use their species-
specific behaviour in situations where it is not needed to obtain a conven-
P. Jensen, F.M. Toat es/AppL A nim. Behav. Sci. 37(1993) 161-181 175

tional reward, or even where it decreases the reward to the animal, is taken as
evidence for the hedonic value of the motor act. For example, in pigeons placed
in a context associated with food delivery, the behaviour of pecking a lighted
key will emerge even if to do so actually delays the arrival of food (see Gard-
ner and Gardner, 1988 ). This may very well mean that behaviour in itself is
reinforcing to the animal, but to establish that behaviour per se is positively
reinforcing, one has to devise an experiment in which the opportunity to be-
have is made contingent upon a response, but for which no extrinsic conse-
quence is derived. This is not an easy task.
The fact that animals will frequently work for food even in the presence of
free food is sometimes taken as a sign of a need to perform feeding behaviour
(reviewed by Gardner and Gardner, 1988 ). Forkman ( 1991 ) found that ger-
bils faced with a choice between digging food out of bowls containing differ-
ent concentrations of grain hidden in sand, would choose to take a large
amount of food out of the bowls which required a lot of digging, even if that
actually reduced the food intake per unit time. Considering only this result,
one might be led to draw the conclusion that the performance of digging has
a hedonic value to the gerbils. However, the result only held true when the
animals were tested in consistent situations, that is, the bowls were always on
the same places and the gerbils knew where to find high and low concentra-
tions of food. When the bowls were swapped around between the trials, the
animals more consistently chose to forage from the high concentration bowls
and so did not use a lot of 'unnecessary' digging movements. While the first
part of the experiment might lead to an interpretation in terms of digging
being reinforcing, the second part suggests no such thing. Rather, the need of
the gerbils may be connected with the consequence of digging, e.g. gaining
information (Forkman, 1991 ). Digging as such, therefore, would not qualify
on a 'global list' of gerbil needs, since it varies with environmental
circumstances.
Other evidence may come from the fact that the performance of certain
behaviours may affect the physiological state of the animal. Dantzer and
Morm6de ( 1981 ) showed that pigs in a frustrating situation had a relatively
lower post-test plasma corticosteroid level if they had the opportunity to pull
a chain during the trial, compared with pigs that had no such possibility. For
rats, Weiss et al. ( 1976 ) found that the possibility to perform aggressive be-
haviour (i.e. only the motions, not actually being engaged in a fight) de-
creased the development of gastric ulcerations following foot shock treat-
ment. This suggests that behaviour may play a role in coping and in that sense
it may of course be rewarding.
The strong tendency of animals in barren situations to develop stereotyp-
ies, i.e. motor patterns repeated over and over again for long periods of time,
is also frequently suggested as a sign o f b e h a v i o u r being rewarding, since it is
often parts of thwarted actions that develop into stereotypies (Hughes and
Duncan, 1988a). Even if this may be the case it certainly does not hold for all
176 e. Jensen, ~: M. Toates /AppL A nim. Behav. Sci. 37 (1993) 161-181

stereotypies in all situations and caution is needed when discussing the signif-
icance of these behaviour abnormalities (Mason, 1991 a,b).
In conclusion, it does appear that the performance ofbehaviour per se may
be reinforcing under some circumstances. The mere action may also be a cop-
ing response in stressful situations (Weiss et al., 1976). However, it remains
to be investigated what effects motor activity may have on internal factors in
specific motivational systems. From the point of understanding the needs of
animals the essential question to answer, concerning specific activities, is the
degree of contribution to the negative feedback that the motor activity in it-
self provides (see Fig. 1 ). Even in simple, specific acts, this is likely to vary
with environmental circumstances. We also cannot see the reason why a be-
haviour should be regarded as a need if the motion patterns are self-reward-
ing, whereas it would not if the activity was indirectly rewarding.

Discussion

We have presented arguments against the 'catalogue approach' to behav-

ioural needs. None of the commonly used criteria, such as the degree of inter-
nal contribution to motivation, time-dependent build-up of tendency, and
the behaviour being self-rewarding, are valid for distinguishing non-needs.
In this paper and in an earlier account (Toates and Jensen, 1991 ) we have
argued that a given behaviour will inextricably be a function of both external
and internal factors. To give general weightings to each is, in our opinion,
conceptually wrong. Although current motivational models usually incorpo-
rate the action of both external and internal factors, each tends to weight one
of them as more important. When attempting to understand the needs of an-
imals, such weighting is misleading.
Maybe the main controversy in theories of motivation concerns the issue
of goal directedness (Toates, 1986; McFarland, 1989). The picture emerging
from the present account is one where goal directedness is an essential part of
the control of the behavioural output. It appears that animals often attempt
to perform at least some parts of their species-specific behaviour patterns re-
gardless of the environment (as in the case of nest building), but without the
assumption that the consequences of the behaviour are compared with a de-
sired goal, most of the findings reviewed in this paper are difficult to explain.
So, in the light of motivation theory, what is a 'behavioural need'? Ob-
viously, the dichotomy between internally and externally controlled behav-
iour is not very useful. Is there any sense at all in splitting the activities of a
certain species into those constituting 'behavioural needs' and other behav-
iour? We cannot see the reason for this. However, this should not be misun-
derstood to mean that animals have no needs with regard to their behaviour.
On the contrary, we have shown evidence that the inability to engage in spe-
P. Jensen, F.M. Toates/AppL Anim, Behav. Sci. 37(1993) 161-181 177

cies-specific behaviour may cause signs of suffering (e.g. abnormal behav-

iour, pathology) and the mere possibility to perform some behaviour may
decrease the physiological effects of stressful situations. What we are arguing
against is the 'catalogue approach', aiming at classifying some specific behav-
iour patterns of a given species as needs, as opposed to other parts of the
behavioural repertoire.
There will presumably always be a need to feed in the sense that harm will
come from a failure to feed. However, even here the distress might also be a
function of context (as discussed earlier). Thwarting of feeding might lead to
frustration and therefore negative welfare consequences, but so might thwart-
ing of sex. Whether one wants to describe behaviour as a need therefore de-
pends upon knowledge of the environmental context. In the absence of aver-
sive stimulation we would not suggest that an animal has a need to display
aggressive behaviour. However, to take an extreme case, as was discussed, if
a rat is subject to aversive stimulation, performing an attack can reduce signs
of stress and contribute to positive welfare consequences. Thus, one might
want to call it a need in this situation.
In considering ethological needs, it is essential to understand the conse-
quences to the animals of not being able to perform a certain behaviour. For
instance, Van Liere and Bokma ( 1987 ) showed that the feathers of hens, de-
prived of dust-bathing for 3 days were less fluffy than those of control ani-
mals. Also, different substrates are not equally effective in obtaining the main
functional effect of the dust-bathing, that is, removal of feather lipids, which
is mirrored in different capacities for thermo-insulation of the plumage (Van
Liere and Siard, 1991 ). Knowing such effects of non-performance o f a behav-
iour provides some possibilities to assess the importance of it to the animal.
This is obtained by focusing on the feedback arrows rather than the causation
arrows in Fig. 1.
Ultimately, what should guide our welfare considerations is the degree of
suffering an animal will experience if a certain behaviour is impaired. This
requires a holistic approach to the causation and control ofbehaviour and we
suggest that it should aim at clarifying the different pathways outlined in Fig.
1 for specific behaviour patterns under consideration. Considering, for ex-
ample, the possible needs involved in the performance of dust-bathing in hens
we should not limit our discussion to the role of internal versus external fac-
tors in the causation, but put equal effort into understanding the feedback
processes that control and eventually shut off the motivation, as well as the
possible goal representations and the way in which differences between these
and the consequences of the behaviour affect the output. When these aspects
are fully understood we will have a good basis for judging what are the welfare
consequences that result if the behaviour is prevented.
Our conclusion is that animals do possess needs with regard to their behav-
iour, i.e. prevention of performance of species-specific behaviour may cause
178 P. Jensen, F.M. Toates/Appl. Anita. Behav. Sci. 37 (1993) 161-181

signs of reduced welfare. However, to assign some particular behaviour pat-

terns as 'behavioural needs' as opposed to others that would not qualify for
this denotation is misleading, in particular when this is based on the assumed
degree of internal control of the behaviour. The needs an animal may have
once a motivational system is activated, be it through the action of internal
or external factors, is a complex of obtaining a goal and performing species-
specific behaviour on the route to the goal. There may probably be needs as-
sociated with all behaviour and the hierarchy of needs will vary according to
environmental situation.

Acknowledgements

We wish to thank Drs Bj/~rn Forkman and Linda Keeling for valuable com-
ments on the manuscript. The work on these ideas was partly done when Dr
Fred Toates was a visiting scientist at the Department of Animal Hygiene in
Skara, financed by a grant from the Swedish Board for Agricultural and For-
estry Research.

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