Professional Documents
Culture Documents
Clinical E t h o l o g y in Food A n i m a l P r a c t i c e
ABSTRACT
Luescher, U.A., Friendship, R.M., Lissemore, K.D. and McKeown, D.B., 1989. Clinical ethology
in food animal practice. Appl. Anim. Behav. Sci., 22: 191-214.
The relationships between ethology and veterinary medicine, and clinical ethology and clinical
veterinary practice, are being discussed. Clinical veterinary ethology is defined as the science
occupied with the diagnosis and treatment of primary behavior disorders, including behavioral
causes for clinical disease and sub-optimal performance. For discussion of behavior problems,
these are categorized into those with direct economic impact, those with indirect economic impact
and those without economic impact. Examples for each category are reviewed. Problems with
direct economic impact are flank biting, tail and ear biting, group aggression, pen fouling and
savaging of piglets by the dam in swine; and non-nutritive sucking, urine drinking, the bu|ler steer
syndrome, milk sucking, and behavior contributing to lesions, teat injury and mastitis in bovids.
Behavior problems with indirect economic impact include restlessness, mutual massaging, copro-
phagia and stereotypies such as bar biting in swine; preputial sucking and urine drinking in swine
and bovids; and object and mutual licking, abnormal motor patterns, extended lying down inten-
sions and tongue rolling in bovids. Abnormal behavior without economic impact is discussed in
terms of welfare implications. Practical recommendations are made concerning the integration of
behavior observations into a herd health program and regarding the solution of behavior problems.
It is concluded that clinical ethology makes a significant contribution to veterinary practice, es-
pecially to herd health programs. More information is needed, though, especially relative to: (1)
the epidemiology of behavior problems; (2) the economic effects of problems and their treatment;
(3) the efficacy of treatments.
INTRODUCTION
a widely accepted branch of veterinary medicine. One of the many reasons for
this might be its more integrative and less analytical character compared to
the more traditional areas of veterinary science such as pharmacology or ge-
netics, etc.
Many scientists have spoken for the integration of applied ethology into the
veterinary teaching program (e.g., Fox, 1967; Houpt, 1976; Voith, 1984; Broom,
1987 ). Today, ethology is included in the curriculum at several veterinary col-
leges (Stricklin, 1983 ) and into various veterinary texts (e.g., Fraser and Mays,
1986; Leman et al., 1986). The following discussion will focus on some of the
reasons for this development.
There is an intimate relationship between physiology and behavior, in that
many of the regulatory mechanisms of an organism necessary to maintain
homeostasis and grant normal development or reproduction, imply behavior
(Tschanz, 1982). Interference with normal behavior can, therefore, also be
expected to hamper important physiological functions. This has indeed been
shown for the courting behavior in boars which plays a significant role in fer-
tility and fecundity of the sow (Hemsworth, 1978), and the periparturient be-
havior of the sow influencing the speed and ease of piglet births (Vestergaard
and Hansen, 1984). A veterinarian needs to know about such complex inter-
actions between behavior and physiology in order to successfully solve cases of
sub-optimal herd performance.
Moreover, behavior has a complex relationship to clinical disease; disease
obviously affects behavior (e.g., Hart, 1985), but the reverse may be true as
well. For example, Madec (1982) suggested that prolonged sitting of sows in
crates may be one reason for frequent urinary tract infection and Sanford
(1982) suggested that impairment of social thermoregulation in pregnant sows
may cause abortions. Some aberrant behavior, such as stereotypic behavior,
must be considered a disease in itself, since it implies severe pathophysiological
changes in the affected animal (Dantzer, 1985). Behavior, in some instances,
can also be an important vector in disease transmission (Visnjakow and Geor-
giev, 1972).
With the development of preventive veterinary medicine and herd health,
knowledge of behavior gains increasing importance. A herd health approach is
performance rather than disease oriented. Many economically important per-
formance traits, however, are directly related to behavior. Fertility of dairy
cows, for instance, depends on the display of estrus behavior (Hurnik, 1987);
Productivity of sows necessitates appropriate maternal behavior (Fraser, 1977);
and rate of gain in turkey poults depends on their ability and motivation to
ingest the provided feed (Lewis and Hurnik, 1979). Impairment of behavior
such as mentioned above obviously results in reduction of performance and,
thus, becomes clinically relevant.
Finally, behavior problems are a frequent cause for veterinary consultations,
especially in companion animals (McKeown and Luescher, 1988), but also in
193
farm animals (Luescher and McKeown, 1987). Their prevalence and impor-
tance require that the practicing veterinarians have a basic knowledge of eth-
ological principles in order to provide a satisfactory service to their clients.
A DESCRIPTION OF CLINICALVETERINARYETHOLOGY
(Wright, 1986) listed only two behavior scientists working in this field, only
one of them being a veterinarian. Although this survey cannot be considered
complete, the number is certainly small.
(3) The individual animal approach is ill-suited for dealing with behavior
problems in farm livestock. At a scientific level, an epidemiological approach
is necessary or desirable (Ewbank, 1986), possibly in tandem with a laboratory
experiment (Webster, 1983 ). At the practical level, problems have to be dealt
with on a herd basis (Luescher and McKeown, 1987).
The apparent lack of reliable data and the as yet limited applicability of
behavior science in food animal practice contributes, as we believe, to the mea-
ger recognition this discipline has among veterinarians. This paper is written
to draw attention to this deficit and to promote the development of clinical
veterinary ethology in food animals.
CATEGORIZATIONOF BEHAVIORPROBLEMS
Most producers are concerned about behavior problems only if they have an
obvious direct economic impact. Such problems are not uncommon and may
be very costly. Nevertheless, for most of them, exact figures regarding preva-
lence and economic significance are missing.
Pigs (usually weanling pigs) have been reported to massage each other's
flank or lower rib cage with vertical movements of the nose (Allison, 1976).
This behavior abnormality is considered to derive from frustrated suckling
behavior (van Putten and Dammers, 1976). Our own observations indicate
that the incidence of flank sucking appears to be significantly greater in piglets
that are weaned before 20 days of age compared to piglets weaned at a later
age. Algers (1984) observed an influence of weaning age on frequency of flank
sucking, but only when piglets were housed in cages. This behavior can escalate
to actual flank biting and persist up to slaughter weight. It often results in
various degrees of distress and damage to the victim (Allison, 1976), and is
associated with reduced growth of both the victims and the instigators (Fraser,
1978). Prevention involves improvement of the climatic environment, wean-
ing the piglets at an older age, weaning them into an enriched environment
(Algers, 1984) and housing the weaner pigs by litter rather than in large groups.
Treatments such as physical separation of pigs, keeping the pigs in the dark,
or even inserting rings into their noses (Allison, 1976), merely treat the symp-
toms and are not satisfactory long-term solutions.
No other behavior problem in farm animals has stimulated both the interest
and ingenuity of researchers as much as tail biting in pigs. A large number of
possible causes have resulted in a similarly large number of suggested treat-
ments, such as manganese supplementation, a higher protein ration and some
as far fetched as spraying the pigs with perfume (von Cramon, 1978).
Tail biting begins as a harmless nibbling of the tail, presumably a re-directed
investigative behavior ( Steiger and Arnold, 1976), and is well tolerated by the
target animal. Only in a later stage, once the tail is injured and blood is drawn,
does it involve rigorous attacks on the tails of the victims (van Putten, 1969 ).
This latter stage may be an expression of predatory aggression and appetite
for blood (Fraser, 1987).
It is known that males are more prone to the condition than females (about
twice as common in males, Penny et al., 1981) and that there is a seasonal
influence on the prevalence of tail biting (Penny and Hill, 1974). Suggested
196
Ear biting is a re-directed activity similar to tail biting, but involves chewing
and biting of the tip or the base of the ears of pen mates. It may be performed
as a stereotypy, alternating with vacuum chewing. In spite of some epidemiol-
ogical differences (Penny and Hill, 1974), ear biting is generally precipitated
by the same factors as tail biting. The incidence appears to have increased since
producers have begun to routinely dock tails. Like tail biting, ear biting may
well have its origin in the nursery and persist later in spite of adequate envi-
ronmental conditions, and possibly straw bedding, in the grower barn. In rare
cases, this behavior can have devastating economic effects due to ear loss and
death of affected pigs. The treatment is the same as for tail biting. Particular
attention should be paid to any factors contributing to ear lesions, which might
in turn trigger ear biting, such as fighting after re-grouping, fighting over the
waterers or ear mange infestation (Straiton, 1967).
(4) Group aggression in fattening pigs towards one member of the group
One pig in the pen, usually neither the smallest nor the largest, is singled
out and then repeatedly attacked either by one pig at a time or by the whole
197
group at once. The victim is quickly exhausted and if not removed in time, dies.
Post-mortem examination reveals no pathological changes other than super-
ficial skin lesions. Once the victim is dead or removed, the group often turns
towards another group member. The condition has been observed in pens with
good footing, often with litter, ample space and rather large groups. The inci-
dences always followed management procedures which induced a high state of
arousal such as cleaning out the pen, catching one pig out of the pen or, in one
case, playing with the pigs. The condition is generally rare and even in affected
herds usually occurs only sporadically, but in some cases the economic loss can
be significant, especially when finisher pigs are involved. Treatment is directed
towards reducing arousal by handling the pigs calmly and by keeping them in
smaller groups.
Pigs are naturally clean animals. Although they are not programmed to
maintain one defecation site, when given the choice they defecate and urinate
away from their lying, feeding and drinking areas. Various factors have been
identified as influencing the pigs' choice of dunging site (Petherick, 1983 ). Pen
fouling occurs when pigs are too crowded, when the ambient temperature is
too high, or when the designs of the pen and/or the ventilation system are
faulty. Solid walls enclosing the lying area seem to be a particularly important
factor. Where fence partitions are used, pen fouling can usually be corrected
by installing solid pen partitions. Some clinical cases indicate that a fully slat-
ted floor and high stocking densities during the growing period can subse-
quently result in pen fouling in partially slatted or solid floor barns during the
finishing period. Weight gain and feed conversion may be adversely affected
by pen fouling (Petherick, 1983) and labor is increased.
but provision of straw did not reduce pre-partum restlessness (Hansen and
Curtis, 1981 ) and was not successful in preventing savaging in practice. Un-
satisfactory rearing conditions of gilts may increase their tendency to savage
(Schouten, 1987). In rare cases, an imbalance in iodine metabolism may be
the underlying cause (Stockl et al., 1970). There are obvious differences in
incidence between different genetic strains. Some gilts seem to be behaviorally
immature for raising piglets when farrowing for the first time.
Control measures include the assessment and correction of stress factors,
improvement of management around farrowing, genetic selection and possibly
breeding the gilts later. The use of farrowing crates with solid sides to allow
for social isolation and to provide a degree of shelter for the sow is a promising
approach. If sows are given the choice, they prefer such crates over conven-
tional ones with bars.
Short-term treatment may include the immediate removal of the piglets dur-
ing parturition and returning them after parturition is over. Treatment with
tranquilizers may be successful (Symoens and van Gestel, 1972 ), but does not
address the causes of the condition.
Veal calves kept on a liquid diet, which is consumed rapidly, suck various
parts of each other's body such as the scrotum, ears and prepuce (Sambraus,
1984), or inanimate objects (Kopp et al., 1986). The amount of sucking de-
pends on early sucking experience (de Wilt, 1985), on the method of feeding
(Szucs et al., 1983 ) and on the presence of effective releasing stimuli for suck-
ing (Mees and Metz, 1983). The behavior may have clinical sequelae (Groth,
1978). It is as yet unclear if it can develop into milk sucking as the animals
grow older (Grommers, 1979). Non-nutritive sucking is interpreted as re-di-
rected sucking (Kopp et al., 1986), resulting from frustration of the sucking
drive (Sambraus, 1984). More frequent meals (Mees and Metz, 1983) or pro-
vision of a pacifier (Kopp et al., 1986) increase the amount of non-nutritive
sucking, while it can be reduced by separating the calves from their dam at
birth and raising them on teat buckets (de Wilt, 1985), extending the feeding
period and increasing the effort involved in sucking by providing milk through
a rubber teat with a small orifice. Separation of the calves after feeding for 10-
20 min greatly reduces sucking on body parts of pen mates (Sambraus, 1984),
but this has to be considered a merely symptomatic treatment.
In large groups of feedlot steers, a few members of the group may attract
others which will repeatedly mount them. The bullers, i.e., the mounted ani-
mals, may show decreased weight gain and even die, if not removed in time.
199
Bullers are also less resistant to respiratory disease (Lincoln, 1983). Predis-
posing factors are conditions which impose stress on the animals, such as stormy
weather, crowding, large and unstable groups, changes in the group (Brower
and Kiracofe, 1978; Irwin et al., 1979) and boredom (Ulbrich, 1981). Other
contributing factors may be unique to the bullers themselves, such as hormonal
or metabolic abnormalities (which may be consequences rather than causes),
their behavior (standing to be mounted), pheromones, different hair color
(Brower and Kiracofe, 1978) or their apparent inability to integrate into the
social hierarchy of the group (Klemm et al., 1983). Implants (Irwin et al.,
1979), particularly if not applied properly (Lincoln, 1983), and phytohor-
mones (Pierson et al., 1976) are considered as causes as well.
The buller steer syndrome may result in significant economic losses. Surveys
in Colorado (Pierson et al., 1976) and Kansas (Brower and Kiracofe, 1978),
indicated that 2.9 and 2.2% of steers became bullers, and the loss per buller
was estimated at $6 and $23, respectively. To prevent the problem, the man-
agement and production system should be evaluated with regard to the above-
mentioned factors and corrected if found deficient. On a short-term basis, bul-
lers may be separated, their implant removed, or their odor masked with strong
smelling substances. Application of progesterone is also mentioned as symp-
tomatic treatment (Lincoln, 1983).
1980; Deja et al., 1982) and devices to be mounted on the cows' nose (Sam-
braus, 1985a) have been developed for symptomatic treatment of the condi-
tion. In light of the probable relationship of milk sucking to inadequate rearing
and housing systems, and its association with tongue rolling, such treatments
must be considered inadequate as long-term solutions.
system, as in the case with poorly designed cubicles (Kammer and Tschanz,
1975; Cermak, 1987 ) or tie stalls (Groth and Eichler, 1978) or from preventing
normal movement patterns or stances, such as species-typical rising or normal
body position when eating (Groth, 1985). Features of the barn environment
inducing such behavior abnormalities and injuries have been reviewed by var-
ious authors, e.g., Kammer and Tschanz, ( 1975 ), Fessl et al. (1984) and Groth
(1984, 1985 ). Analysis of the problem involves inspection of each individual
cow, assessing the physical properties of the housing system and behavior
observations.
This list of behavior problems with direct economic impact in farm animals
is by no means exhaustive. The few and tentative indications of economic sig-
nificance and recommendations for treatment reflect the scarcity of applicable
research in this area and the lack of clinical experience pertaining to the cor-
rection of these problems. For more information on how to practically ap-
proach a primary behavior problem, the reader is referred below to Practical
Considerations.
DEFICIENCIES
IN
ENVR
I ONMENT,
~#lJ MANAGEMENT ~
ABNORMAL ~ POORHEALTH
BEHAVIOR = AND PERFORMANCE
(DISEASE)
Fig. 1. The relationship between production system, behavior and performance.
202
(4) Bar biting, vacuum chewing and other stereotypies in pregnant sows
(5) Preputial sucking and urine drinking in fattening pigs and beef cattle
Abnormal ways of rising, lying down and atypical lying positions are com-
monly due to physical restriction when veal calves are housed in crates (Bog-
ner and Reissig, 1982; Webster et al., 1985 ). A normal resting position in ster-
nal recumbancy may be important for various physiological functions (van
Putten and Elshof, 1982; de Wilt, 1985).
A high frequency and long duration of lying down intensions in dairy cows
(Andreae et al., 1982) and a higher than normal proportion of time spent
standing while ruminating can be interpreted as signs of poor adaptation to
the housing system and stress.
This stereotypic behavior has been described in calves (Unshelm et al., 1982;
Wiepkema et al., 1987), fattening bulls (Sambraus and Gotthard, 1985) and
dairy cattle (Sambraus, 1985a). It is considered to be a vacuum feeding behav-
ior caused by a lack of roughage (Unshelm et al., 1982; Sambraus, 1985a). It
is one of few stereotypic behaviors displayed by ruminants (Sambraus, 1985b ).
The indirect economic impact of abnormal behavior is poorly documented.
Thus, the validity of figures on the economic impact of behavior problems such
as the ones available for tail biting in pigs or for the buller steer syndrome,
must be doubted since usually only direct costs are considered (i.e., mortality,
carcass devaluation, etc.). One approach to demonstrate the indirect costs of
behavior is to consider the reduction in productivity of the animal performing
a given behavior, not just the one suffering from its consequences, as was done
for belly nosing in piglets (Fraser, 1978) and for stereotypies in tethered sows
(Cronin et al., 1986). However, those animals within a herd which perform
abnormal behavior may actually be better off than their herd mates not per-
forming it (Wiepkema et al., 1987; McGlone and Blecha, 1987). A correct as-
205
BEHAVIORPROBLEMS WITHOUTAPPARENTECONOMICIMPACT
PRACTICALCONSIDERATIONS
(B) Observations
(C) Measurements
Specific measurements relating to the physical, the climatic and the social
environment are taken.
A comparison between the natural situation and the artificial barn environ-
ment helps to identify shortcomings of the latter, such as the lack of suitable
target objects for behavior and the impossibility for the animals to seek social
contact or social isolation.
CONCLUSIONS
ular, to herd health programs. Ethologists have established a solid base of use-
ful concepts on which a clinical approach to behavior problems can be devel-
oped. Since it is such a young discipline, clinical veterinary ethology suffers
from an obvious lack of much needed information which must be established
if it is to become more viable. This information relates mainly to: (1) the epi-
demiology of behavior problems; (2) economic aspects, i.e. direct and indirect
costs of problems and feasibility of treatment regimes; (3) the efficacy of var-
ious treatment regimes to establish accurate prognoses. It is hoped that this
paper will generate interest in the area, and stimulate the research and clinical
ethological practice necessary to provide this information.
REFERENCES
Algers, B., 1984. Early weaning and cage rearing of piglets: Influence on behavior. Zentralbl. Vet-
erinaermed. Reihe, A, 31: 14-24.
Allison, C.J., 1976. Snout-rubbing as a vice in weaned pigs. Vet. Rec., 98: 254-255.
Andreae, U. and Papendieck, T., 1971. Verhalten von Milchkuhen bei der Wahl ihrer Liegeboxen
im Laufstall. Tierzuchter, 23: 432-435.
Andreae, U., Thielscher, H.H., Unshelm, J. and Smidt, D., 1982. Ethological and physiological
conditioning of young cattle in intensive housing systems. In: J.P. Signoret {Editor), Welfare
and Husbandry of Calves. Martinus Nijhoff, Boston, pp. 235-241.
Appleby, M.C. and Lawrence, A.B., 1987. Food restriction as a cause of stereotypic behavior in
tethered gilts. Anim. Prod., 45: 103-110.
Bengtsson, A.C., Svendsen, J. and Andersson, M., 1984. Housing of sows in gestation: studies of
behavior in different types of group housing. In: J. Unshelm, G. van Putten and K. Zeeb (Ed-
itors), Proceedings of the International Congress on Applied Ethology in Farm Animals, 1984,
at Kiel, pp. 214-216.
Blackshaw, J.K. and McVeigh, J.F., 1984. Stereotype behavior in sows and gilts housed in stalls,
tethers, and groups. In: M.W. Fox and L.D. Mickley (Editors), Advances in Animal Welfare
Science. pp. 163-173.
Blood, D.C., Morris, R.S., Williamson, N.B., Cannon, C.M. and Cannon, R.M., 1978. A health
program for commercial dairy herds. 1. Objectives and methods. Aust. Vet. J., 54: 207-215.
Bogner, H. and Reissig, F., 1982. Verhalten yon Mastkalbern in verschiedenen Haltungssystemen.
Tierarztl. Prax., 10: 465-470.
Broom, D.M., 1987. The veterinary relevance of farm animal ethology. Vet. Rec., 121: 400-402.
Brower, G.R. and Kiracofe, G.H., 1978. Factors associated with the buller-steer syndrome. J.
Anim. Sci., 46: 26-31.
Brummer, H., 1978. Verhaltensstorungen. In: H.H. Sambraus (Editor), Nutztierethologie. Paul
Parey, Berlin/Hamburg, pp. 281-292.
Buchenauer, D., 1981. Parameters for assessing welfare, ethological criteria. In: W. Sybesma (Ed-
itor), The Welfare of Pigs. Martinus Nijhoff, Boston, pp. 75-89.
Burmeister, F., Teuffert, J. and Schluter, H., 1981. Die bedeutung des Milchsaugens fur die Eu-
tergesundheit. Monatsh. Veterinaermed., 36: 407-411.
Cermak, J., 1987. The design of cubicles for British Friesian dairy cows with reference to body
weight and dimensions, spatial behavior and upper leg lameness. In: H.K. Wierenga and D.J.
Peterse (Editors), Cattle housing Systems, Lameness and Behavior. Martinus Nijhoff, Bos-
ton, pp. 119-128.
210
Craig, J.V., 1981. Domestic Animal Behavior. Prentice-Hall, Englewood Cliffs, NJ, 364 pp.
Cronin, G.M., 1985. The Development and Significance of Abnormal Stereotyped Behaviors in
Tethered Sows. Thesis, Wageningen, pp. 19-50.
Cronin, G.M. and Wiepkeman, P.R., 1984. An analysis of stereotyped behaviour in tethered sows.
Ann. Rech. Vet., 15: 263-270.
Cronin, G.M., van Tartwijk, J.M.F.M. van der Hel, W. and Verstegen, M.W.A., 1986. The influ-
ence of degree of adaptation to tether housing by sows in relation to behavior and energy
metabolism. Anim. Prod., 42: 257-268.
Dantzer, R., 1986. Behavioral, physiological and functional aspects of stereotyped behavior: a
review and reinterpretation. J. Anim. Sci., 62: 1776-1786.
Dantzer, R. and Mormede, P., 1983. De-arousal properties of stereotyped behavior: Evidence from
pituitary-adrenal correlates in pigs. Appl. Anim. Ethol., 10: 233-244.
Davies, G., 1985. Art, science and mathematics: New approaches to animal health problems in the
agricultural industry. Vet. Rec., 117: 263-267.
Deja, D., Partzsch, C. and Vedder, H., 1982. Erste Erfahrungen beim Einsatz der "Frohndorfer
Methode" gegen das Saugen der Rinder im Bezirk Erfurt. Monatsh. Veterinaermed., 37: 132-
134.
De Wilt, J.E., 1985. Behavior and welfare of veal calves. Thesis, Wageningen, 138 pp.
Drawer, K., 1973. Concrements and pseudo-concrements in food animals. Vet. Med. Rev., 1973:
160-166.
Ekesbo, I., 1966. Disease incidence in tied and loose housed dairy cattle. Acta Agric. Scand., Suppl.
15, 74 pp.
Ekesbo, 1978. Ethics, ethology and animal health in modern Swedish livestock production. In:
D.W. Fiilsch (Editor). The Ethology and Ethics of Farm Animal Production. Birkhauser,
Basel/Stuttgart, pp. 46-50.
Ewbank, R., 1973. The trouble with being a farm animal. N e w Sci.,60: 172-173.
Ewbank, R., 1986. Animal Welfare: is an epidemiological approach possible? In: M.V. Thrusfield
(Editor), Proceedings of the Society of Veterinary Epidemiology and Preventive Medicine, 2-
4 April 1986, at Edinburgh, Gt. Britain, pp. 92-96.
Fessl, L., Hantak, E. and Hofmann, R., 1984. Zur Problematik des Baues von Rinderstallungen
aus orthopadischer Sicht. Berl. Muench. Tieraerztl.Wochenschr., 97: 235-239.
Fox, M.W., 1967. The place and future of animal behavior studies in veterinary medicine. J. Am.
Vet. Med. Assoc., 151: 609-615.
Fraser, A.F., 1984. Some divisions of applied ethology. Appl. Anita. Behav. Sci.,12: 201-207.
Fraser, A.F., 1985. Deprivation of maintenance behavior in modern farm animal husbandry. In:
A.F. Fraser (Editor), Ethology of Farm Animals. Elsevier,Amsterdam, pp. 377-389.
Fraser, C.M. and Mays, A. (Editors), 1986. Merck Veterinary Manual. Merck and Co., Rahway,
NJ, 1677 pp.
Fraser, D., 1975. The effectof straw on the behavior of sows in tether stalls.Anim. Prod., 21: 59-
68.
Fraser, D., 1977. Some behavioural aspects of milk ejection failureby sows. Br. Vet. J., 153: 126-
133.
Fraser, D., 1978. Observations on the behavioural development of suckling and early-weaned pig-
letsduring the firstsix weeks afterbirth. Anita. Behav., 26: 22-30.
Fraser, D., 1987. Attraction to blood as a factor in tailbiting by pigs. Appl. Anita. Behav. Sci.,17:
61-68.
Gonyou, H.W., Hemsworth, P.H. and Barnett, J.L., 1986. Effects of frequent interactions with
humans on growing pigs. Appl. Anita. Behav. Sci., 16: 269-278.
Graf, B., Wegmann, R. and Rist, M., 1976. Das Verhalten von Mastkalbern bei verschiedenen
Haltungsformen. Schweiz. Landwirtsch. Monateh., 54: 333-355.
Grommets, F.J., 1979. A review of the problem of milk-sucking in dairy cows. Appl. Anim. Ethol.,
5: 293.
211
Groth, W., 1978. Tierschutz und verhaltensbezogene Gesichtspunkte der Kalbermast. Tierzucher,
10: 419-422.
Groth, W., 1984. Mangel im Haltungssystem als Ursache von Gliedmassenschaden bei Rindern.
Tieraerztl. Umsch., 39: 196-201.
Groth, W., 1985. Kriterien fur die Beurteilung von Haltungssystemen fur Milchkuhe und Mast-
bullen aus Klinischer Sicht. Tieraerztl. Umsch., 40: 739-750.
Groth, W. and Eichler, H.J., 1978. Haltungsbedingte Schaden beim Milchvieh. Fortschr. Veteri-
naermed., 28: 34-43.
Hansen, K.E. and Curtis, S.E., 1981. Prepartal activity of sows in stall or pen. J. Anim. Sci., 51:
456-460.
Hart, B.L., 1985. Animal behavior and the fever response: theoretical considerations. J. Am. Vet.
Med. Assoc., 187: 998-1001.
Hemsworth, P.H., 1978. The importance of stimulation by partners on the reproductive success
of pigs. International World Congress on Ethology Applied to Zootechnics, 1978, at Madrid.
Houpt, K.A., 1976. Animal behavior as a subject for veterinary students. Cornell Vet., 66: 73-81.
Hurnik, J.F., 1987. Sexual behavior of female domestic mammals. In: E.O. Price (Editor), Farm
Animal Behavior. The Veterinary Clinics of North America, Food Animal Practice, 3. Saun-
ders, Toronto, pp. 423-461.
Irwin, M.R., Melendy, D.R., Amoss, M.S. and Hutcheson, D.P., 1979. Roles of predisposing fac-
tors and gonadal hormones in the buller syndrome of feedlot steers. J. Vet. Med. Assoc., 174:
367-370.
Jongebreur, A.A. and Smits, A.C., 1982. Group housing systems for veal calves. In: J.P. Signoret
(Editor), Welfare and Husbandry of Calves. Martinus Nijhoff, Boston, pp. 217-225.
Kammer, P. and Tschanz, B., 1975. Untersuchungen zur tiergerechten Haltung von Milchvieh in
Boxenlaufstallen. Schweiz. Landwirtsch. Forsch., 14: 203-223.
Klastrup, 0., Bakken, G., Bramley, J. and Bushnell, R., 1987. Environmental influences on bovine
mastitis. Bulletin of the International Dairy Federation, 217, 37 pp.
Klemm, W.R., Sherry, C.J., Schake, L.M. and Sis, R.F., 1983. Homosexual behavior in feedlot
steers: an aggression hypothesis. Appl. Anim. Ethol., 11:187-195.
Kopp, M.B., Friend, T.H. and Dellmeier, G.R., 1986. Effect of feeding method on nonnutritive
oral activities in holstein calves. J. Dairy Sci., 69: 3094-3099.
Leman, A.D., Straw, B., Glock, R.D., Mengeling, W.L., Penny, R.H.C. and Scholl, E., (Editors),
1986. Diseases of Swine, sixth edition. Iowa State University Press, Ames, IA, 930 pp.
Lewis, C.J. and Oakley, G.A., 1970. Treatment of puerperal psychosis in sows with sedative and
anaesthetic drugs. Vet. Rec., 87: 616-617.
Lewis, N.J. and Hurnik, J.F., 1979. Stimulation of feeding in neonatal turkeys by flashing lights.
Appl. Anim. Ethol., 5: 161-171.
Lincoln, S.D., 1983. Ethology and control of buller syndrome. Norden News, 58: 24-26, 28.
Littlejohn, A., 1969. An approach to clinical veterinary ethology. Br. Vet. J., 125: 46-48.
Luescher, U.A. and McKeown, D.B., 1987. Clinical veterinary ethology in food animals. Can. Vet.
J., 28: 304-305.
Madec, F., 1982. Urinary problems in the sow. Pig Int., 12: 28-29.
McGlone, J.J. and Blecha, F., 1987. An examination of behavioral, immunological and productive
traits in four management systems for sows and piglets. Appl. Anim. Behav. Sci., 18: 269-286.
McKeown, D.B. and Luescher, U.A., 1988. A case for companion animal behavior in the veterinary
practice. Can. Vet. J., 29: 74-75.
Meek, A.J., Mitchell, W.R., Curtis, R.A. and Coate, J.F., 1975. A proposed information manage-
ment and disease monitoring system for dairy herds. Can. Vet. J., 16: 329-340.
Mees, A.M.F. and Metz, J.H.M., 1983. Saugverhalten von Kalbern - - Bedurfnis und Bedriedigung
bei verschiedenen Trankesystemen. Tagung Dt. vet. med. Ges.e.V., Fachgruppe Verhaltensfor-
schung, 16-19 November, Freiburg, i. Br.
212
Mellinger, R., 1980. Chirurgie de la langue chez le bovin teteur. Point Vet., 10: 11-14.
Penny, R., 1982. Tail biting most common cause still unknown. Int. Pigletter, pp. 1-2.
Penny, R.H.C. and Hill, F.W.G., 1974. Observations of some conditions in pigs at the abbatoir
with particular reference to tail biting. Vet. Rec., 94: 174-180.
Penny, R.H.C., Waiters, J.R. and Tredget, S.J., 1981. Tail biting in pigs: A sex frequency between
boars and gilts. Vet. Rec., 198: 35.
Peterse, D.J., Rutgers, B., Schaftenaar, W. and Grommers, F.J., 1978. Een onderzoek naar melk-
zuigen by runderen. Tijdschr. Diergeneeskd., 103: 485-489.
Petherick, J.C., 1983. A biological basis for the design of space in livestock housing. In: S.H.
Baxter, M.R. Baxter and J.A.D. MacCormack (Editors), Farm Animal Housing and Welfare.
Martinus Nijhoff, Boston, 343 pp.
Pierson, R.E., Jensen, R., Braddy, P.M., Horton, D.P. and Christie, R.M., 1976. Bulling among
yearling feedlot steers. J. Am. Vet. Med. Assoc., 169: 521-523.
Rushen, J.P., 1984. Stereotyped behaviour, adjunctive drinking and the feeding periods of teth-
ered sows. Anim. Behav., 32: 1059-1067.
Rushen, J.P., 1985. Stereotypies, aggression and the feeding schedules of tethered sows. Appl.
Anim. Behav. Sci., 14: 137-147.
Sambraus, H.H., 1979. Analmassage und Kotfressen bei Mastschweinen. Dtsch. Tieraerztl. Woch-
enschr., 86: 45-84.
Sambraus, H.H., 1981. Abnormal behavior as an indicator of immaterial suffering. Int. J. Stud.
Anim. Probl., 2: 245-248.
Sambraus, H.H., 1984. Gegenseitiges Besaugen yon Kalbern bei kunstlicher Aufzucht. Berl.
Muench. Tieraerztl. Wochenschr., 97:119-123.
Sambraus, H.H., 1985a. Mouth-based anomalous syndromes. In: A. Fraser (Editor), Ethology of
Farm Animals. Elsevier, Amsterdam, pp. 391-422.
Sambraus, H.H., 1985b. Stereotypies. In: A.F. Fraser (Editor), Ethology of Farm Animals. Else-
vier, Amsterdam, pp. 431-441.
Sambraus, H.H. and Gotthard, A., 1985. Praeputiumsaugen und Zungenspielen bei intensiv ge-
haltenen Mastbullen. Dtsch. Tieraerztl. Wochenschr., 92: 465-468.
Sambraus, H.H. Kirchner, M. and Graf, B., 1984. Verhaltensstorungen bei intensiv gehatenen
Mastbullen. Dtsch. Tieraerztl. Wochenschr., 91: 56-60.
Sanford, S.E., 1982. Fall abortions in sows. Can. Vet. J., 23: 36.
Schluter, H., Teuffert, J. and Burmeister, F., 1981. Untersuchungen zum Saugverhalten, zur Hau-
figkeit und zu den Ursachen des Milchsaugens. Monatsh. Veterinaermed., 36: 403-407.
Schmidt, G.H. and Pritchard, D.E., 1987. Effect of increased production per cow on economic
returns. J. Dairy Sci., 70: 2695-2704.
Schouten, W.G.P., 1987. Effects of rearing conditions on the behavior of gilts around farrowing.
Appl. Anim. Behav. Sci., 17:367 (abstract).
Schunke, B., 1980. Verhaltensanomalien bei Zuchtsauen im Kastenstand. Thesis, Munich, 126
pp.
Signoret, J.P., Baldwin, B.A., Fraser, D. and Hafez, E.S.E., 1975. The behavior of swine. In: E.S.E.
Hafez (Editor), The Behavior of Domestic Animals. Bailliere Tindall, London, pp. 295-329.
Simonsen, H.B., 1983. Vurdering af maelkeran og tungerulning pa basis af sporgeskemaunderso-
gelse i 24 problembesaetninger. Dan. Veterinaertidsskr., 66: 669-671.
Smith, W.J. and Penny, R.H.C., 1986. Behavioral problems including vices and cannibalism. In:
A.D. Leman, B. Straw, R.D. Glock, W.L. Mengeling, R.H.C. Penny and E. Scholl (Editors),
Diseases of Swine. Iowa State University press, Ames, IA.
Steiger, A. and Arnold, J., 1976. Untersuchungen zum Schwanzbeissen bei Mastschweinen. In:
Probleme Tiergerechter Haltung. KTBL, Darmstadt, pp. 56-72.
213
Stein, T.E., 1986. Marketing health management to food animal enterprises. Part II. The structure
of herd health management services. Compend. Cont. Educ. Pract. Vet., 8: $331-$336.
Stockl, W., Bamberg, E. and Enengel, H., 1970. Uber den Jodstoffwechsel bei Schweinen mit
einem Beitrag uber das Ferkelfressen. Wien. Tieraerztl. Monatsschr., 57: 274-279.
Straiton, E.C., 1967. How to Recognize and Treat Common Pig Ailments. Farming Press, Ipswich,
160 pp.
Stricklin, W.R., 1983. A survey of animal behavior-related research and teaching activities in
North American Agricultural and Veterinary Medical Colleges. Int. J. Stud. Anim. Prob., 4:
279-283.
Symoens, J. and van Gestel, J., 1972. Die Behandlung des Ferkelfressens mit Azaperon. Tieraerztl.
Umsch., 27: 170-172.
Szucs, E., Molnar, I., Weber, A., Szollosi, I. and Kishonti, L., 1983. The effects of feeding milk
from nipple-pails or buckets in calf rearing. Acta Agron. Acad. Sci. Hung., 32: 273-284.
Tschanz, B., 1982. Verhalten, Bedarfsdeckung und Schadenvermeidung bei Tieren. Tagung Nutz-
tierkommission Schweizer Tierschutz, Internationale GeseUschaft fur Nutztierhaltung, 23 April
1982, at Bern, Switzerland.
Ulbrich, R., 1981. The buller steer syndrome. Int. J. Stud. Anim. Probl., 2: 261-268.
Unshelm, J. Andreae, U. and Smidt, D., 1982. Behavioral and physiological studies on rearing
calves and veal calves. In: J.P. Signoret (Editor), Welfare and Husbandry of Calves. Martinus
Nijhoff, Boston, pp. 70-76.
Van Putten, G., 1969. An investigation into tail biting among fattening pigs. Br. Vet. J., 125:511-
517.
Van Putten, G., 1983. Bewertung von Schweinehartungssystemen bezuglich Wohlbefinden. 3. GFT-
Seminar, 26 September-1 October, Grub.
Van Putten, G. and Dammers, J., 1976. A comparative study of the well-being of piglets reared
conventionally and in cages. Appl. Anim. Ethol., 2: 339-356.
Van Putten G. and Elshof, W.Y., 1982. The lying behavior of veal calves up to 220 kg. In: Signoret,
J.P. (Editor), Welfare and Husbandry of Calves. Martinus Nijhoff, Boston, pp. 83-97.
Vestergaard, K. and Hansen, L.L., 1984. Tethered versus loose sows: ethological observations and
measures of productivity. I. Ethological observations during pregnancy and farrowing. Ann.
Rech. Vet., 15: 245-256.
Visnjakow, J.I. and Georgiev, M., 1972. Swine caudophagy, a new epizootiological link of trichi-
nellosis in the industrial swine farms. Acta Parasitol. Pol., 20: 597-604.
Voith, V.L., 1984. Why should veterinarians study animal behavior? Mod. Vet. Pract., 65: 363-
364.
Von Cramon, A., 1978. Kannibalismus-- ein ungelostes Problem? Dtsch. Tierarztl. Wochenschr.,
85: 345-380.
Webster, A.J.F., 1983. Environmental stress and the physiology, performance and health of rum-
inants. J. Anim. Sci., 57: 1584-1593.
Webster, A.J.F., Saville, C., Church, B.M., Gnanasakthy, A. and Moss, R., 1985. The effect of
different rearing systems on the development of calf behavior. Br. Vet. J., 141: 249-264.
Wiepkema, P.R., 1983. On the significance of ethological criteria for the assessment of animal
welfare. In: D. Smidt (Editor), Indicators Relevant to Farm Animal Welfare. Martinus Nijhoff,
Boston, pp. 71-79.
Wiepkema, P.R., 1985. Abnormal behaviors in farm, animals: ethological implications. Neth. J.
Zool., 35: 279-299.
Wiepkema, P.R., van Hellemond, K.K., Roessingh, P. and Romberg, H., 1987. Behavior and
abomasal damage in individual veal calves. Appl. Anim. Behav. Sci., 18: 257-268.
Wierenga, H.K., 1983. The influence of the space for walking and lying in a cubicle system on the
behavior of dairy cattle. In: S.H. Baxter, M.R. Baxter and J.A.C. MacCormack (Editors),
Farm Animal Housing and Welfare. Martinus Nijhoff, Boston, pp. 171-180.
214
Wood-Gush, D.G.M., Duncan, I.J.H. and Fraser, D., 1975. Social stress and welfare problems in
agricultural animals. In: E.S.E. Hafez (Editor), The Behavior of Domestic Animals. Baillere
Tindall, London, pp. 182-200.
Wright, J.C., 1986. Directory of animal behavior consultants. Animal Behavior Consultant News-
letter, October, 1986.