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Journal of Feline Medicine and Surgery (2018) 20, 411–421

CLINICAL review

Normal feliNe behaviour

... and why problem
behaviours develop
John Bradshaw

Introduction Practical relevance: Cats are

descended from a solitary, territorial
Clinicians have been describing and classifying cats’ problematic ancestor, and while domestication has
behaviours for several decades, but everyday relationships between reduced their inherited tendency to be
cats and their owners remain remarkably antagonistic towards all animals larger than
undocumented. As a species, domestic cats their typical prey, they still place more reliance on
Everyday
have been the subject of many behavioural the security of their territory than on psychological
relationships and ecological studies,1 but often as proxies attachments to people or other cats, the exact
for other, less accessible carnivores, or, more opposite to dogs. Many feline problem behaviours
between cats stem from perceived threats to this security,
recently, because conservationists perceive
and their owners their predatory instincts as damaging to often due to conflicts with other cats. Others are
wildlife,2 so the relevance of these studies to more developmental in origin, often caused by
remain remarkably clinical practice can only be inferential. inadequate exposure to crucial stimuli, especially
Because they live literally under the noses of people, during the socialisation period. Strongly
undocumented. aversive events experienced at any age can also
their doting owners, it is easy to imagine that
the behaviour of pet cats presents no myster- contribute. A third category comprises normal
ies; but, until recently, science has paid very behaviour that owners deem unacceptable,
little attention to how cats relate to the very such as scratching of furniture.
surroundings in which most behavioural dis- Evidence base: This review identifies three areas
orders become apparent – the home, and its in which basic research is inadequate to support
human and animal occupants. Classification widely employed concepts and practices in feline
and diagnosis of these disorders has therefore behavioural medicine. First, classification of cats’
come to be based upon a mixture of: problem behaviours relies heavily on approaches
< Comparisons with other species (most practitioners treat more derived from studies of their behavioural ecology
dogs than cats); and, to some extent, extrapolation from canine
< Custom and conjecture (as there has been so little published studies. Few studies have focused on cats in the
research, most writings on feline behaviour problems are home, the environment in which most behavioural
unavoidably a rehash of what has gone disorders are expressed. Secondly, cats’ chemical
before); senses (olfactory and vomeronasal) are far more
< Common-sense extrapolations sensitive than our own, making it difficult for owners
SERIES OUTLINE or clinicians to fully comprehend the sensory
from what is known about
This article forms part of a series information upon which they base their behaviour.
feline cognition and cat–cat of evidence-based reviews on feline
social behaviour. it is this last Thirdly, although the concept of psychological
behaviour, independently written by key
category that forms the basis distress is widely invoked as an intervening variable
opinion leaders, spanning three Special
of this review. in behavioural disorders, there are still no reliable
Issues of JFMS. An outline of the series
measures of distress for pet cats in the home.
is included as supplementary material
alongside the article at: cpsi.jfms.com Global importance: Psychological distress of
some kind is the primary cause of many of the
behavioural problems presented to clinicians, but
surveys indicate that many more cats display the
same clinical signs without their owners ever
seeking help. The welfare of this ‘invisible’ group
could be improved by veterinarians taking a more
John Bradshaw proactive approach to educating their clients about
BA(Oxon), PhD(Soton) the behavioural needs of pet cats.
University of Bristol Veterinary School, UK
Email: j.w.s.bradshaw@bristol.ac.uk

doi: 10.1177/1098612X18771203
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specimens from around the world,9 doubling

Feline behaviour: some fundamental misunderstandings the archaeologists’ estimate of the time since
Cats are often portrayed as easier to keep than dogs, providing little incentive domestication began, from 5000 to 10,000 years
for owners to consider how best to provide for their newly acquired pet. ago, towards the end of the Neolithic period.
Fundamental misunderstandings of feline behaviour and motivation abound. The first phase, for which it must be said
For example, unlike dogs, which use play as a form of social transaction, adult there is little supporting archaeological evi-
cats’ motivations for ‘playing’ are focused around predatory behaviour, during dence, appears to have been essentially evolu-
which they become largely oblivious to their owner’s involvement (see box tionary, as a small number of wildcats found
below). Many owners believe that their cat’s behaviour is driven by emotions themselves capable of adapting to the new
such as jealousy and pride, and cognitive abilities such as deliberate decep- niche created by the concentrations of rodent
tion, none of which are compatible with biologists’ current understanding of prey around the poorly protected food stores
the feline brain. A substantial minority believe that their cats are incapable of our hunter-gatherer ancestors. With the
of feeling anxious, the emotional response that is implicated in problematic advent of cereal-growing, this niche would
behaviour more than perhaps any other.7 Such misconceptions lead directly have become yet more profitable, permitting a
to the development of problem behaviours: new owners given basic infor- genetic separation between those wildcats
mation on what to expect of their cats, and how to manage their environ- that occasionally raided farmsteads but other-
ment, report far fewer problems than owners left to find out for themselves.8 wise retained their normal hunting behaviour,
and those few cats that happened to have the
right temperament to live permanently within
human settlements, thereby generating the
New owners given basic information on what
new subspecies Felis silvestris catus that even-
to expect of their cats, and how to manage their tually became the pet cat of today. Tolerance
of humans would have been essential to allow
environment, report far fewer problem behaviours these cats to exploit both the prey and the
than owners left to find out for themselves. shelter (crucially, denning sites) available
within these early villages. At some point
within the first 1000 years or so of this devel-
The process of domestication – oping relationship, our ancestors began to
and its shaping of feline behaviour transport cats from place to place, as shown
by the appearance of cats on Cyprus (which
The generally accepted framework for under- has no native wildcat) about 9000 years ago.
standing the normal behaviour of the house- To begin with, these cats would probably
hold cat is by extrapolation from its wild have been almost as solitary and territorial as
ancestors and feral counterparts, rather than their wild counterparts, a tendency that per-
direct observation of behaviour in the home sists, albeit in a diluted form, in our pet cats.
(in stark contrast to human psychology, in However, with the emergence of the first
which the evolutionary approach is a relative- towns, the quantity of vermin available in a
ly new player and still pretty controversial). small area would have increased to more than
our understanding of the derivation of the could feed a pair of cats and their offspring,
domestic cat from its ancestral species, the and tolerance of the proximity of other cats
African/Arabian wildcat Felis silvestris lybica, would have become an important adaptation,
has been revolutionised by recent analyses of gradually evolving into the cooperative social
the dNA of both living cats and museum behaviour seen in today’s farm cat colonies.10

Behaviour we label as ‘play’

In cats, the behaviour that we label as ‘play’ is invariably focused ‘toy’: cats play with mouse-sized toys as if they were mice, rat-sized
on some kind of object, whether that be a commercially produced toys as if they were rats. Hunger causes play to become more
toy, a leaf blowing in the wind, or (less advisably) the owner’s fingers intense,4,5 and emboldens cats to interact with large toys that they
or toes. In contrast to dogs, where many of these would normally be wary of, in just the same way
‘games’ have a social purpose,3 the way in which Cats may look like that a hungry cat will double its efforts to achieve
cats play with objects is precisely what would be
predicted if those objects were potential items of
they are playing, ashould kill. If play was a purely social activity, it
be suppressed, not activated, by hunger.
prey.4,5 Fur- or feather-covered exteriors are preferred but in their heads This is not to say that cats do not enjoy play-
over other textures, as are toys with ‘legs’, and also ing with toys. Each segment of the predatory
toys that disintegrate during play, convincing the they are hunting. sequence, from stalking through to close-
cat that it is making progress towards a ‘kill’.4,5 quarters interaction with the prey, is rewarding
Apparently spontaneous movements of the object activate and in itself, whether or not the cat’s efforts eventually result in
intensify play, irrespective of whether there is a human agent consumption of food. For cats that readily engage in play, brief
involved. The movements of the cat’s limbs and jaws precisely mimic ‘games’ can be used as positive reinforcement, providing a use-
those made towards prey that is the same size and shape as the ful alternative to food and petting when training them.6

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Figure 1 Mutual grooming different subspecies, Felis silvestris silvestris in

appears to reinforce bonds
within existing social
Europe and the desert wildcat Felis silvestris
groups. From ‘Cat Sense’,11 ornata in what is now Pakistan and western
used with permission of the
artist, Alan Peters
india. The dNA of today’s pet cats shows little if
any trace of either,9 although there were descen-
dants of indian desert cats living in Turkey in
the seventh century,12 and it is conceivable
that some of their descendants remain as part
of the local pet population, even today. The
slender oriental body plan that subsequently
gave rise to the Siamese and similar breeds
evolved in genetic isolation13 (wild Felis sil-
vestris does not occur in the Far East) and is an
adaptation to humid environments; its dNA
does not point to descent from the superficial-
ly similar desert wildcat, as was once thought.
in Europe, the second millennium wit-
The social organisation of the cat colonies nessed intermittent persecution of cats and
that form around abundant food sources has their owners, slowing the process of domesti-
proved of considerable interest to biologists, cation; pet cats remained a rarity, perpetuating
because so few other species in the cat family live genotypes that maintained wild-type hunting
in permanent groups, the prime exceptions and reproductive behaviour. As a conse-
being the lion and the cheetah. Unlike both of quence, the behavioural differences between
these, males of the domestic cat do not cooperate The trend today’s pet cats and their wild ancestors may
with one another; it is the females that form be traceable to changes in as few as a dozen
the nucleus of social groups, in which related towards the genes.14 it was not until the 19th century that
individuals assist one another in the raising of keeping a cat primarily for its company
kittens. The behaviours that knit these groups cat becoming became at all widespread in Western Europe
together – mutual flank-rubbing and groom- the most or the United States, and the trend towards
ing (Figure 1) – have obvious parallels in the cats becoming the most popular pet in both
popular pet
A second factor that undoubtedly slowed
way that many pet cats behave towards their regions only took off a few decades ago.

domestication is the cat’s hypercarnivory. All

owners,10 including rubbing around their legs
in Western
felids require a flesh-based diet, due to a set of
and attempting to lick their hands, for example.

ancient mutations that restricted their ability to

The second phase, domestication proper, Europe and the
process plant-derived foods. In addition to
began in Egypt about 5000 years ago, which is
United States
their need for relatively large amounts of sulfur-
where we have the first archaeological evi-

containing amino acids (methionine, cysteine,

dence for cats as pets. it is unclear whether
only took off a
taurine) and the vitamins niacin and thiamine
and to what extent this marked another

in their diet, they also require essential fatty

change in the cat’s behavioural genetics, this few decades
acids of animal origin in order to synthesise
time towards people, or a cultural shift in how
ago.
prostaglandins, which most other mammals
cats were perceived – pet dogs, the obvious

can make from plant-derived precursors. Cats

precedent, had already been popular for

also need a constant supply of high quality

many centuries. Subsequently, cats achieved

protein, as they are unable to switch off amino

considerable religious significance in Egypt,

acid catabolism, again tying them to their car-

before being transported all over the known

nivorous lifestyle. In this sense they are quite

world during the Classical period, beginning

unlike dogs, which can switch to plant-based

around 2500 years ago. The first cats to arrive

foods when meat is scarce. The cat’s basic

in Britain were probably brought there by

nutritional requirements were not properly

Phoenician traders, about 300 BCE. For the

understood until the 1970s, so it has only been

next 1000 years or so most cats would have

since then that cats have been able to repro-

been pets second and mousers first. Unlike

duce successfully without supplementing

dogs, which would have been most helpful

their diet through hunting. This provides one

when trained to perform whatever tasks were

explanation for their persistent habit of

required of them, cats control vermin most

patrolling and attempting to defend a hunting

effectively when left to their own devices,

territory, even when well fed by their owners:

counteracting any tendency for cats to become

too few generations have passed since this has

ever more sociable as they adapted to living at

no longer been a necessity for their fundamen-

increasingly higher densities.

tal motivations to have changed appreciably

during the Graeco-Roman period, domestic

(although in theory these might be reduced in

cats were transported both east and west, into

the future by directed breeding).15

areas where they came into contact with and
could interbreed with native wildcats of

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Biological factors influencing problem behaviours in cats

< Unlike dogs, derived from the highly social wolf, domestic < During their domestication, cats evolved a simple, female-based,
cats are descended from a solitary territorial ancestor. social system, but their default is still to be antagonistic towards
< Until the latter half of the 20th century, cats were mainly unfamiliar cats (dogs have lost the wolf’s natural aggression
valued for their abilities as independent hunters, for which towards members of other packs). Multi-cat households
a social bond with humans or other cats was unnecessary. are usually artificial assemblages of unrelated individuals,
< Cats have specialist nutritional requirements that impeded and therefore chronic antagonism is likely
their becoming wholly dependent on their owners. between at least some pairs of individuals.
< Cats prioritise territory over social interaction with their Cats residing in adjacent households are
owners, the opposite to dogs. unlikely to interact amicably.

Cats’ sexual behaviour is also largely Cats as individuals –

unchanged from that of their wild ancestors. learning and ‘personality’
Pedigree cats being still in the minority, the
majority of pet cats are the offspring of Although cats are generally (though
unplanned matings that occur when a female wrongly) perceived as being less train-
comes into season, attracts one or more entire able than dogs, learning plays a major
males and engages in wild-type courtship part in the way both species behave.
behaviour that originally evolved to ensure max- Thus how a given cat reacts to a par-
imum fitness for her offspring. (Nowadays in the ticular situation will vary considerably
West, the critical factor determining a cat’s life- depending on its lifetime experience;
time reproductive success, both male and female, albeit genetic variation between indi-
must be to escape being neutered,15 but the full viduals does also play a part, if only
effects of this change, which should lead to those in channelling learning in different
cats that are most difficult to socialise produc- directions. The environment that cats
ing the most offspring, have not yet permeat- find themselves in, and its match
ed through to the behaviour of the typical cat.) with their formative experiences, also
By comparison with dogs and other common appears to play a critical role.20
domestic mammals, cats are not yet fully domes- The most important determinant of
ticated. They select their own mates, and retain how a cat reacts to humans is its expe-
much of the predatory drive of their wild ances- riences during the socialisation period,
tors, both of which require interventions from which in this species runs from 2–9
their owners to counteract their effects. There weeks of age, earlier than in the dog.
Figure 2 Gentle handling promotes
is an increasing trend, especially in urban
socialisation in kittens of all ages. From Kittens that have not been handled by
areas of the United States and in continental
‘Cat Sense’,11 used with permission of the the age of 9 weeks do not approach
artist, Alan Peters
Europe, to keep cats indoors throughout their people spontaneously, and usually
lives, denying them the opportunity to mate, become ‘feral’, the most likely scenario being that
presuming they are not neutered, and also to their mother was also feral and therefore chose
hunt wildlife, in addition to protecting them to give birth and nurse her kittens in some out-
from hazards such as diseases and road traffic. of-the-way location. This is not to say that 9-
The cost of such confinement to their mental week-old kittens have learned everything about
wellbeing has been much debated but inade- people that they ever will; rather, handling dur-
quately quantified.16,17 in the UK, cats allowed ing the socialisation period opens a window that
outdoors are frequently neutered18 and may enables continued positive interaction with
be fitted with a bell or other deterrent device humans and promotes learning of how to inter-
to reduce their impact on bird and small mam- act with them (Figure 2). This process seems
mal populations, although the most effective to be particularly formative during the next
suppressor of ‘serious’ hunting behaviour is 4 months or so, such that for most cats the style
probably a nutritionally complete diet,19 such in which they interact with people becomes
as most pet cats in the West receive. fixed by the time they are about a year old.21,22
The corresponding optimum period for learn-
ing how to interact with other cats has not been
studied systematically, which is unfortunate,
By comparison with dogs, cats are not yet fully since incompatibilities between cats in the same
domesticated. They select their own mates, and or neighbouring houses cause a significant pro-
portion of problem behaviours. Hand-raised
retain much of the predatory drive of their wild kittens that do not receive any contact with
ancestors, both of which require interventions from other cats during the socialisation (to humans)
period are anecdotally considered to be at
their owners to counteract their effects. enhanced risk of developing problems such as

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nervousness, aggression and a reduced ability For most cats The subjective world of the cat –
to cope with changes in their environment. senses
others may become overattached to their rais- the style in
ers or subsequent owners, attempting to suck- Cats being mammals, it is easy for owners to
le from fingers or elbows even when adult. which they presume that the world that they themselves
The vast majority of cats that are presented at interact with perceive is identical to their cat’s. The biological
clinics for behavioural problems are neither feral reality is that subjective impressions of their
nor hand-raised, but nonetheless display consid- people surroundings differ between our two species
erable variation in ‘personality’ from one indi- at three levels: the information gathered by
vidual to another. Pedigree cats as a whole do
becomes fixed the sense organs, the manner in which this
appear to be more likely to be presented with by the time information is integrated and filtered by the
problem behaviours. However, this may be brain, and the emotional reactions that are
due as much to owner expectations and emo- they are about triggered. Were more owners to fully compre-
tional and financial investment as to differences hend these differences, and react appropriately,
a year old.
in the cats’ experiences – ordinary domestic many problem behaviours might not escalate
cats or ‘moggies’ are typically raised informally to the point of requiring veterinary intervention.
in family surroundings and rehomed at 8 Cats’ sense of hearing and vision are both
weeks, whereas pedigree cats are often bred in different to our own, even though the corre-
purpose-built housing and are normally homed sponding sense organs are constructed along
at 12–16 weeks of age, potentially restricting the same mammalian pattern.30 The range of
their socialisation. There are undoubtedly frequencies that cats can detect encompasses
genetically based differences in temperament all of those that we can hear, except for very low-
between breeds,23 but there is little hard data pitched notes, which cats can probably detect
on how these translate into the prevalence of using the sensitive pads on their toes. Cats can
behavioural disorders. also hear more than an octave above our own
Non-pedigree pet cats also vary in tempera- range, into the ‘ultrasound’ region: this is an
ment and therefore in their susceptibility to dif- adaptation that enables cats to eavesdrop on
ferent problems. Early attempts to classify the the ultrasonic calls that small rodents use for
main personality types suggested that there are communication. in terms of the physics of how
three broad categories: cats that are confident the mammalian ear works, this ability is unre-
and inquisitive, cats that appear timid and ner- markable, as the hearing range of the ear
vous, and cats of both kinds that are more or less should go up as it gets smaller. What is actu-
active than the norm. The bold/shy axis, which ally unexpected is the cat’s ability to hear low-
has also been detected in many other species, can pitched sounds, including men’s voices. This
have a genetic basis in cats, although some ele- has been traced to a septum that alters the res-
ments are undoubtedly learned.24 Problematic onant properties of the middle ear, which is
behaviour is presumably affected by many also found in other species of small cat, so did
genes; one of these, coding for an oxytocin recep- not evolve during domestication; rather, it was
tor, has been recently identified as contributing a lucky accident that enabled even the earliest
to irritability.25 More such identifications will domestic cats to detect and react to male voices.
doubtless follow, offering the possibility that The cat’s eyes are adapted to provide accu-
genetic typing may eventually become a tool rate vision in very low light. They are large by
in the veterinary behaviourist’s armoury. comparison with our own, with even larger
Although the assessment of temperament pupils, both factors maximising the amount of
differences between cats has recently received light entering the eye. The reflective tapetum
attention from scientists,26 this has mostly been behind the retina further increases the effi-
based on owners’ subjective perceptions of their ciency of the eye (and also gives the cat the
cats, rather than measuring their behaviour eye-shine that inspired the eponymous road-
directly:27,28 this method interposes an unhelpful markings). Most of the detectors on the retina
curtain of anthropomorphism between the are rods, with relatively few of the cones that
data gathered and genuine and reproducible give us our daytime colour vision; cats are not
characteristics of the cats themselves. A more only red-green colour blind, but when their
objectively based tool – the Feline Behav- brain interprets the information coming from
ioural Assessment & Research Questionnaire the retina, it prioritises outline, brightness and
(Fe-BARQ) – has recently been validated, and especially movement over colour, almost the
has demonstrated the complexity of cat-to-cat exact opposite to ourselves.
variation in behaviour, identifying no fewer than
17 underlying types of variation, including play- Cats being mammals, it is easy for owners
fulness, sociability to people, attention-seeking
and fear of other cats.29 it may eventually pro- to presume that the world that they themselves
vide clinicians with a useful starting point for perceive is identical to their cat’s.
assessment of their patients’ basic ‘personalities’.

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each several experiences of odour collected

from the other, well before any face-to-face
encounter.6 Seemingly, what each animal
learns from this process is that the cat bearing
the unfamiliar odour is in the vicinity (how
else would its smell have got there?) but has
not presented any threat. When the other cat
is eventually introduced, the resident cat will
Disturbances be able recognise it by its odour (and vice
of the olfactory versa) and both will be less likely to default
to defensive mode. other pre-exposures to
environment unfamiliar scents in neutral or deliberately
rewarded contexts presumably work through
make a a combination of habituation and positive
significant association: examples include habituation to
the odour of items brought into the house in
contribution preparation for a new baby. olfactory enrich-
ment is effective for cats in rehoming cen-
to so many tres,31 and therefore probably for pet cats also.
Figure 3 Flehmen. From ‘Cat Sense’,11 used with permission of problem Although domestic cats routinely deposit
the artist, Alan Peters scent marks on objects in their environment,
behaviours remarkably little is known about their func-
of cats. tion.32 The external ears, the temples, the
Hearing and vision are sufficiently similar cheeks and the corners of the mouth all pro-
between cats and their owners that there are at duce scents, some or all of which must be
least common reference points: this is not true deposited when a cat cheek-rubs an object or
of olfaction. Akin to most mammals, with the another cat, and there is also a gland beneath
exception of higher primates like ourselves, the chin that is discharged during chin-
cats rely on odour both to communicate with rubbing behaviour (Figure 4). Scent-
one another and to gather information about producing glands are additionally found
many aspects of their environment. Not only between the toes, which presumably leave
are their noses about a thousand times more scent behind whenever the cat scratches with
sensitive than ours, cats possess a second its claws. There is another cluster of odour-
olfactory detection system, the vomeronasal producing structures at the rear end, includ-
organ. Lying between the hard palate and the ing the preputial and anal glands, the scent
nostrils, this acts as a kind of half-way house from which can be dispersed by urination and
between smell and taste, mainly detecting defecation, respectively, as well as skin glands
chemicals that have dissolved in saliva as the in front of and down the length of the tail.
cat opens its mouth in the characteristic
grimace known as ‘gaping’ or ‘Flehmen’
(Figure 3). Reliance on these chemical senses
is so removed from our own experience that it
is difficult enough for biologists and veteri-
narians, never mind owners, to imagine what
the world must be like for an animal that is far
less visually obsessed than we are. it is there-
fore unsurprising that what we presume
(unable to experience them for ourselves) are
disturbances of the olfactory environment
make a significant contribution to so many
problem behaviours of cats.
Although there has been very little research
into how cats interpret the odours they
encounter, the importance they place on them
is indicated by the effectiveness of behaviour
modifications using odour cues. Thus, for
example, a cat that has lost its characteristic
odour due to a period of hospitalisation may
be attacked by other cats in the household
when it returns. Anecdotally, the process of
introducing two previously unacquainted cats
to one another can be smoothed by allowing Figure 4 Main scent-producing structures of the domestic cat

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Cats’ brains have relatively The subjective world of the cat –

cognition and emotion
large areas devoted to olfaction,
Great strides have been made over the past
and also balance, as befits two decades in understanding the cognitive
their lifestyle. abilities of dogs; in addition to testing their
behaviour, it has proved possible to train dogs
to undergo fMRi (functional magnetic reso-
nance imaging), which has revealed, for
it is claimed – though the primary data example, signs of emotional responses specific
appears only in a patent application – that to images of the dog’s owner. Cats, though
the various facial glands of domestic cats equally trainable,6 lag far behind in corre-
produce several chemical compounds (or sponding research.26 However, it has been
mixtures) that have distinct behavioural possible to show that cats do form genuine
effects.33,34 one named F3 is widely used in attachment bonds with their owners, and
behaviour modification in its commercial modify their behaviour accordingly.21,22 Thus,
form (Feliway; Ceva). it is stated that this despite the scepticism of some, there is a logi-
mimics a chemical signal that cats deposit cal basis for separation disorders in (perhaps a
around their familiar home ranges and ‘helps small minority of) cats.
in organising the environment by classifying Some inferences can be made about feline
it into known objects and unknown cognition based upon what we now know
objects’.33,34 about dogs, since their brains are both con-
From an adaptationist perspective this structed according to the carnivore pattern,
seems an unlikely hypothesis. Mammalian and are therefore substantially different to our
chemical signals are formally divided into own. Relative to the size of their bodies, cats’
two types: pheromones, which have a brains are less than half the size of ours, and
communicative function common to all much of the difference is due to our compara-
individuals within a species, and ‘signature tively huge cerebral cortex, the ‘thinking’ part
mixtures’ that identify each individual to of our brains. Cats’ brains have relatively
others (and quite possibly to the individual large areas devoted to olfaction, and also
that deposited the scent upon returning to balance (the cerebellum), as befits their
later to the same site).35,36 Pheromones are lifestyle. in terms of behaviour, the structure
deployed in situations where it is advanta- of their brains suggests that cats almost cer-
geous to both the emitter and the receiver tainly live much more in the present than we
that some simple message is understood, do, neither ruminating on the past nor plan-
and so vary little between individuals; the ning for the future.
odour given off by a female in season is a These cognitive differences also have con-
case in point (note F3 is marketed as a sequences for cats’ emotional repertoires.
‘pheromone’). Signature mixtures vary from Contrary to what many owners believe, cats
one individual to another and have to are probably incapable of experiencing emo-
be learned by recipients. The anal gland tions such as guilt, pride and grief, all of
secretions of cats fall into the category of which require a sense of self and/or a concept
signature mixtures,37 differing reproducibly of past, present and future. (Cats do appear to
from one cat to another, and it seems likely grieve for missing feline or human compan-
that this is also true of the cheek gland ions, but this behaviour can be neatly
secretions. explained by the lingering odour of the absen-
Since it is implausible that any artificial tee, undetectable to us but all too real to the
odour could match that of the cat in cat.) Simpler emotions, what we might refer to
whose house it was being dispersed, the as ‘gut feelings’, such as anger, affection, fear
cat should logically interpret the synthetic and anxiety, are generated in the limbic sys-
odour as belonging to some other cat that tem, which is common to all mammals. due to
has invisibly scent-marked its territory. the differences between their brains and ours,
Lack of clarity as to the mode of action of cats may not experience these quite in the way
Feliway may explain why different inves- we do, but it is difficult to explain cat
tigators have found very different levels of behaviour without invoking the triggering of
efficacy.38,39 these simpler emotional states.

It is difficult to explain cat behaviour without invoking the triggering

of simpler emotional states, such as anger, affection, fear and anxiety.

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Behavioural disorders – why A change in such as ‘spite’ to the cat when a change in its
‘problem’ or unwanted behaviour environment causes it to urinate elsewhere in
develops their cat’s the house. Under these circumstances, pun-
behaviour is ishing the cat for its supposed transgression is
A change in their cat’s behaviour is often what likely to lead to a vicious spiral in which the
prompts owners to consult a veterinarian, often what anxiety that caused the original incident
whether the cause of that change is ultimately becomes magnified at every recurrence.
psychological (a ‘problem behaviour’) or prompts Resolution of house-soiling requires compre-
physiological (eg, a pathology or a hormonal owners to hension of the cat’s perspectives, including a
disorder). it is conventional to divide location that smells very slightly of urine and
behavioural problems of cats into those that consult a faeces (to the cat’s nose, and probably un-
are essentially adaptive responses insofar as noticeable to the owner), thus indicating a
the cat is concerned, but inconvenient or dis-
veterinarian, location that has been used previously; but
tressing for the owner, and those that have whether the equally, not strongly, which might otherwise
medical causes, such as epilepsy or hyperthy- indicate an area that has been over-used and
roidism; although, as has been pointed out,40 cause of that would in the wild present a risk of infection
in real life many cases comprise elements of change is by parasites.42 in multi-cat households, indi-
both, and variable combinations of adverse vidual cats that tend to avoid one another will
early life experiences and psychological stress ultimately be especially wary while urinating and defe-
can lead to or exacerbate medical conditions, cating, but very often such cats will not have
such as chronic lower urinary tract signs.41 psychological been provided with multiple litter trays.
The focus of the remainder of this discussion or physiological. As a second example, many owners are
is exclusively behaviour that can be interpret- annoyed by their cat’s instinctive habit of
ed as a normal adaptive response to the situa- scratching on friable surfaces, behaviour that
tion the cat finds itself in. is perfectly natural from the cat’s perspective,
owners’ lack of understanding of normal and only a problem to the owner. Scratching
cat behaviour can lead to errors in manage- can be redirected away from furniture and
ment which, in turn, impact negatively on curtains using standard training techniques,
welfare. To take one common example, house- yet declawing (onychectomy) is still widely
soiling; many owners do not appreciate that used, where legal, as a less labour-intensive
cats that use indoor litter trays rely on cues option. However, in multi-cat households
such as location, odour and texture of the sub- declawing doubles the risk of house-soiling,
strate when choosing where to eliminate. The possibly because these cats are less able to
cat may not ‘know’ that the litter box is the defend themselves against other cats, so
place that the owner wants them to urinate potentially replacing one problem with anoth-
and defecate, but some owners evidently er while at the same time threatening the cat’s
believe the opposite, attributing motivations welfare.43,44

Owner psychology vs cat welfare: a mismatch

There appears to be a considerable mismatch
between the occurrence of potentially problem-
atic behaviour in the pet cat population, and its
prevalence in terms of numbers of enquiries to
advice services and cases presented to practi-
tioners (Figure 5). Behaviour that causes incon-
venience or embarrassment to owners, especially
urination and defecation in undesired locations,
is over-represented in clinical surveys; behaviour
that may be perceived as ‘just what cats do’,
such as aggression between cats living in neigh-
bouring households, is under-represented, even
though these may be different expressions of the
same underlying issue. In one door-to-door survey,
Figure 5 Percentages of pet cats reported to display problematic behaviour. Data were
over half of all owners reported that their cat was gathered from a door-to-door survey of owners (UK), spontaneous enquiries to an ‘ask the cat
fearful of unfamiliar people and/or neighbourhood expert’ website advertised on the Discovery Channel (USA), and referrals of cats to the UK-
based Association of Pet Behaviour Counsellors (APBC). Mutil./pica/anxiety/attn. = total for
cats,45 yet it seems that few owners perceive
self-mutilation, pica, separation anxiety and attention-seeking45
such behaviour as a problem worth addressing,
regardless of its effect on their cat. Thus although owners, and their cat’s best interests at heart, their conception of what con-
especially those who seek professional help, are likely to have stitutes good welfare in their pet may be significantly skewed.46

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Most behavioural disorders seem to result

from the cat being prevented from achieving KEY poiNts
emotional equilibrium, either due to a per-
ceived or actual external threat, or frustration < The conceptual basis for the
at being prevented from performing its natu- classification of cats’ behavioural
ral behaviour. Such cats are conceived of as (non-medical) disorders is much less
experiencing distress,47 which then leads to secure than a quarter-century of
changes in behaviour such as enhanced generally concordant literature might
aggression or flight behaviour, or attempts to suggest.
< There has been very little research
manipulate the olfactory environment by
changing the location of scent-marking,
into the behaviour of cats in ‘normal’
including urination and defecation. Cats may
households (ie, those that do not
perceive threat when their access to an ade-
seek advice from clinicians), but it
quate home range is or becomes restricted,
seems likely that in many of these
including being kept indoors,48 changes in
the owners are simply tolerant of
household routine, or evidence that other cats
behaviour that others consider
or other animals are intruding into the cat’s
problematic. The population of
core areas: changes in odour profile must play
owners, and thus cats, seen by
a large, possibly predominant role in all of
clinicians may therefore select
these scenarios. More direct social stressors
itself more on the basis of owner
include physical conflict with other cats, both
psychology than cat welfare.
within and outside the owner’s house, and
with humans. The latter may be due to the cat < The widespread assumption that
having adopted inadequate or malfunctional many disorders are due to general
strategies for reacting to certain people or, psychological distress has not been
indeed, all humans, usually a consequence of proven. Explanations will have to be
inadequate socialisation, or having received found as to why some cats respond
aversive experiences at the hands of its own- in one way, others in
ers, such as physical punishment.47 another; why some cats
While perceived threat may be conceptually seem to have their own
useful in diagnosing unwanted behaviour, it ways of coping, at least to
has not proved straightforward to derive reli- their owners’ satisfaction,
able independent measures of distress in pet while others do not.
cats. For example, the stress hypothesis pre-
dicts that multi-cat households should contain
a higher proportion of stressed individuals
than among single cats; however, three recent
independent studies have been unable to con- Acknowledgements
firm this.49–51 The Cat-Stress-Score, a compos-
ite behavioural measure validated for use on The author wishes to thank Tony Buffington, Mikel
caged cats, is not easy to apply in domestic Maria delgado, Emily Blackwell and two anony-
situations.49 Physiological methods might be mous referees for helpful comments on the
more useful for clinicians, but cortisol, the manuscript.
hormone that is widely used to assess chronic
stress in other species, seems to be difficult to Conflict of interest
interpret in cats, whether measured in faeces
or in urine. Chronic distress can induce a The author declared no potential conflicts of inter-
range of non-specific signs, including vomit- est with respect to the research, authorship, and/or
ing, diarrhoea, anorexia, fever, lethargy, publication of this article.
feigned sleep and inhibition of grooming or
overgrooming.20 Funding

The author received no financial support for the

research, authorship, and/or publication of this
Most behavioural disorders seem to result from article.
the cat being prevented from achieving emotional
Dedication
equilibrium, either due to a perceived or actual
This review is dedicated to the memory of Sir
external threat, or frustration at being prevented
Patrick Bateson, who did so much to promote the
from performing its natural behaviour. study of the behaviour of domestic cats as a legiti-
mate branch of ethology.

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