AND OTHER SPECIAL SENSES
quite near-sighted and the eyes are reasonably
fixed in position (requiring head-turning to look
around) (Raleigh, Scott, Jackson and Jackson, 1976).
They are dichromats and can see colour but their colour
vision is limited (Beaver, 1992). They can see very well
in dim light as they have a high proportion of rods (which
have a low threshold for activation by light energy) in the
retina. Catsí eyes have tapetum lucidum behind the retina
to reflect light and aid in night vision (Beaver, 1992).
of the catís eye can change in aperture,
opening from about 1 cm in diameter down to a narrow
slit that is barely visible. In the dark the catís pupil
opens much wider than ours and collects
the light so it can see objects we are
unable to (Hart, 1980a). The catís ability
to change pupil size is very important as an adaptation
that allows it to hunt at night (Beaver, 1992). Cats have
a large binocular vision, 100-130į, and a panoramic
field of 250-280į.
are fully open at about day 17 of life
(Beaver, 1992). However, vision is the last sense to
develop fully (Thorne, 1992), needing postnatal time to
mature (Beaver, 1992). The light blink reflex, which
develops at about day 50 of gestation to day 13 of life,
disappears around day 21, due to the development of
acute pupil control (Beaver, 1992).
acute hearing and can hear sounds in the
range of 10-60 KHz (human is in the range of 18-20
KHz), and can produce ultrasonic emissions. The catís
outer ear can be directed towards the source of sound
(Hart, 1980a). The ear pinna can rotate about 180į
(Beaver, 1992) and, by using both ears, cats can accurately
locate sound (Raleigh et al., 1976).
(1992) states that a catís ability to hear
ultrasonic sounds is probably an adaptation to hunting
small prey, such as rats, that emit them, as cats are not
known to make those sounds. Queens and kittens may
use ultrasound as an identifying contact call. More work
is needed to determine the extent of ultrasonic sound
emission (Bradshaw 1992).
ability to hear very small tone differences
at high frequencies and at low frequencies decreases
with age, especially at higher frequencies (Beaver,
1992). The efficiency of the catís middle ear decreases
at frequencies above 10 KHz due to mass limitations in
the ossicles (Bradshaw, 1992).
becomes fully developed by about four
weeks of age (Thorne, 1992).
has a much more acute sense of smell than
we have, and in addition to having a larger olfactory system,
they also have the vomeronasal organ (Jacobsonís
organ). It is thought that the flehmen reaction (curling of
the upper lip) is a behaviour that exposes the
vomeronasal organ to sex pheromones present in the
female catís urine or vaginal area (Hart, 1980a).
is most frequently displayed by tomcats
are used for identification and communication
in adult cats (Thorne, 1992). Scents are also used
to explore and habituate new environments (Beaver,
of smell is highly developed at birth, and
this is very important as it guides the kitten to the mammary
gland for nursing (Beaver, 1992). Kittens use
smell to orientate themselves and recognise home, littermates
and mother. Adults use it to mark their home
territory (with urine) and when meeting (Beaver, 1992).
Cats that lose their sense of smell because of a viral
infection, also lose their appetites, change their toileting
habits, and donít indulge in courtship (Bradshaw, 1992).
ORGANISATION AND DOMINANCE HIERARCHIES
loners and avoid interactions with other cats,
except when with a mate, with young, or if several
belong to the one household. The area travelled during
normal activities is known as the home range (Beaver,
1992. Bradshaw, 1992. Thorne, 1992). It is much larger
for males than for females (Bradshaw, 1992). The range
may overlap other animalsí ranges (Thorne, 1992).
Studies on free-ranging cats (Fox, 1975), showed that
cats have a home territory and a home range that consists
of places for resting, sunbathing and watching.
These places are connected by a network of paths and
are visited regularly. In a neighbourhood, cats have an
order of dominance, which depends on time and place.
If a low-ranking cat has already entered a narrow passageway
and a high-ranking cat enters, the less dominant
animal will sit and wait until the way is clear. Cats
go to great lengths to avoid meeting another cat on a
pathway, and chance face-to-face encounters lead to
fighting and chasing and the development of a dominant
Ėsubordinate relationship. Subordinate males can
be pushed around in a dominant maleís home range
and essentially become nomads (Liberg, 1981, cited in
group of cats is maintained in colony pens, they
should be provided with shelves so they can Ďowní one
and retreat there from other cats (Hart, 1980a). The cats
will work out an arrangement where certain ones use
the floor at different times to others. Rubbing may help
reinforce social positions, with subordinate individuals
generally rubbing more dominant conspecifics
(Macdonald, Apps, Carr and Kerby, 1987, cited in
socialisation period is the time when all primary
social bonds are formed and is the most important period
during the catís life (Beaver, 1992). Active social contact
with more than one adult cat at some crucial development
stage is necessary for an adult cat to adapt later
to social living conditions (Bradshaw, 1992).
social contact is between a female and her
and the motherís behaviour is known as
epimeletic (care-giving) behaviour (Beaver, 1992). A
kitten will also display etepimeletic (care-seeking)
behaviour when in a strange environment (Beaver,
or furniture scratching leaves a visible
and, at the same time, foot gland secretions
give the scratched object a scent that can be detected
by an intruder (Ewer, 1963).
behaviour may be used as a form of stretching
and cats are most likely to do it after waking (Beaver,
1992). The longer an object serves as a scratching
medium, the more significant it is to the cat (Beaver,
1992) because it will represent greater territorial
importance and the cat will have invested
more in its territory.
Spraying (scent marking with urine):
of backing up, raising the tail, which trembles, and
spraying urine, usually on a vertical object. It is most
usually done by tomcats and objects that are sprayed
often tend to be located along territorial boundaries. The
urine marks left can be identified by other cats, so track
can be kept of their neighbours (Hart, 1980a). Spraying
serves to bring the male and female together during the
breeding season, and is commonly done at a height
convenient for sniffing (Beaver, 1992).
Cheek and head rubbing: A
cat often rubs its head or cheek against a
chair, a table or a personís leg. We tend
to think the cat is being friendly, but since Prescott
(quoted in Fox, 1975) found scent glands along the catís
tail, on each side of the catís forehead and on the lips
and chin region, it appears likely that the cat is performing
a type of scent marking of its special territory. This
behaviour is important as a form of tactile communication
in social groups. It also serves a purpose of reinforcing
social positions (Thorne, 1992).
interesting point about the catís marking behaviour,
compared with the dogís, is that a cat marks to
warn other animals to stay away, but a dog marks to tell
others he is there so they can join him.
and tail are areas of the body more commonly
used for cat to cat rubbing (Bradshaw, 1992).
cats, marking is more usually done in front of
conspecifics than when the cat is isolated, perhaps
because it functions as a display of dominance
is normally helpful in stretching muscles
human interaction it may be an indication of
excitement (Thorne, 1992).
can also be performed by any cat in a
stressful situation (such as when there is a new cat in
the house) (Beaver, 1992).
rubbing against a human seems more common
once the catís presence has been acknowledged
(Beaver,1992). This reflects the use of head rubbing
between familiar cats.
or more of the waking time of some cats is
spent in grooming, which includes face washing,
scratching the hair, coat and skin with the claws, licking
the fur and pulling at the claws with the teeth. It is
suggested that cats turn to grooming when
frustrated in an action; this may reduce
important function of grooming is to maintain
healthy skin. It also removes parasites and dander,
and relieves tension (Beaver, 1992).
socially groom if kept in social groups,
such as when there is more than one cat in a household.
(Turner and Bateson, 2000).
communicate with body postures and facial
expressions (Fox, 1974).`There are three basic categories:
offensive threat, defensive threat and a passive
crouched posture. The offensive threat is a stare, with
body poised to attack; the defensive threat is spectacular,
with the back arched, fur fluffed up and the tail
straight up with the fur fluffed. The cat approaches the
enemy sideways with prancing steps ó the sideways
approach and raised fur make the cat look bigger and
fiercer; the passive crouch with tail down is a passive
submission display to appease an
mother and kittens may have an ultrasonic call
that acts as a contact call (Bradshaw, 1992).
communication is especially important to
cat species with solitary habits (Bradshaw, 1992).
communicate using smell (such as spraying)
and leaving visual signs (such as scratching)
communication (rubbing) may aid scent
marking (Thorne, 1992).
communication is important, especially
when the cats cannot see each other, such as when it is
dark, when they are separated (such as two cats on
their own home territory), or when blind newborn kittens
cry to attract their motherís attention (Bradshaw, 1992).
communication includes a range of call
types. Purring is in response to pleasurable contact,
developed as a kitten when full of milk and resting with
mother and littermates, which continues to adulthood,
such as when petted by a human (Bradshaw, 1992). An
inaudible purr is common in human presence
(Remmers and Gautier, 1972).
is for greeting, the growl and yowl for
aggression and the hiss and spit is a defensive reaction.
Sudden, sharp pain may result in the pain shriek
is generally directed at humans (Turner
and Bateson, 2000) and is rarely observed in inter-felid
cat is polyoestrus during the breeding season
and shows distinct oestrous behaviour. She is more
active and nervous than usual and has a loud mating
call to attract males. Ovulation is stimulated by copulation.
The male approaches an oestrous female from
behind or sideways as she assumes a receptive crouch,
elevates the pelvic region and holds her tail to one side.
She also treads with the back legs. The male mounts
and thrusts and the female gives a copulatory cry, and
as soon as the penis is withdrawn the female becomes
aggressive towards the male. There is a display of
postcopulatory behaviour as she rolls and
rubs on the floor (Hart, 1980a; Fox,
assume the receptive crouch (lordosis)
before the male mounts; it is accentuated when the
male grasps her scruff (Thorne, 1992).
will display lordosis again several minutes
after the initial display. The cycle of events will then
be repeated and the repetitions can continue for up to
two days (Bradshaw, 1992).
contact by the male during proestrus is
not tolerated by the female (Beaver, 1992).
is stimulated by copulation, and one
female may mate with a number of males in one
period (Thorne, 1992).
not form pair bonds and the male leaves after
copulation. When birth is close, the cat retires to a dark
quiet place. When the kittens are born and the mother
has licked them to stimulate respiration and cleaned
them, she rests in a semi-circle around them. If a kitten
crawls away she touches or licks it to bring it back.
Nursing starts within an hour or two after birth and for
the first two days, the mother remains constantly with
the kittens. She also initiates nursing and a teat order
develops among the kittens (Ewer, 1961) which reduces
competition. Defecation and urination is stimulated by
the mother licking the anogenital region. She keeps the
nest clean by eating the waste, grooms the kittens and
begins to play with them. By the fifth week, nursing time
declines and the mother begins to teach the kittens
predatory behaviour. The relationship between predatory
and play behaviours has been studied (Caro, 1981)
and it seems that predation develops later than social
play; although some of the patterns are similar, others
increase in development (e.g. chasing and being
chased, or biting), while other behaviours decrease
(e.g., decrease in the number of sequences containing
three kittens). Fostering is easy and mother cats will
readily adopt other young.
her kittens chewing or biting her, the queen
will initially growl at them. If this does not work she may
hit the kittens on the nose, drag them away or turn and
move away from them.
behaviour is the primary social pattern
exhibited by female cats, Kittens spend
almost all their time with their mother or
siblings for their first three weeks of life (Beaver, 1992
& Turner et al. 2000).
often licks and awakens her kittens to
stimulate them to begin suckling (Beaver, 1992).
presence of littermates reduces the stress of
new environments (Bradshaw, 1992).
be early-season oestrus synchrony (Liberg,
1981, cited in Turner and Bateson, 2000) resulting
in litters of a few females being born around the
same time. This would allow communal caring. Feldman
(1993, cited in Turner and Bateson, 2000) found that a
queen involved in communal care would move her kittens
more frequently. This may be because it would be
safer per move than if she was alone, as the kittens at
either site (where being moved from or to) would be
guarded most of the time.
catnip (a member of the mint family) response
(Hart, 1980a) involves the leaves or extract of the plant.
Between 50-70% of cats respond by sniffing, licking or
chewing the material and head shaking and gazing into
space are common. Some cats rub their cheeks and
chin over the catnip, others paw or dig it. The response
lasts for 5 to 15 minutes; it is triggered by the active
ingredient nepetalactone and is mediated through the
catnip response does not seem to be related to
a sexual response or hunting/aggressive behaviour
a strong stimulus to stop purring
response is inherited on a dominant autosomal
gene (Bradshaw, 1992).
AND PROBLEM BEHAVIOURS
weíll try to cover most of the common behaviour
problems. This is just an overview and the extent to
which one can discuss individual therapies is limited.
pet displays behaviours that are unwelcome,
they are described as behaviour problems. What
vets and behaviour therapists have to consider is to
what extent these behaviours are the normal responses
to early social experiences, management and training.
More important, perhaps, we have to distinguish
changes in behaviour that have their origins in disease
since to leave disease untreated while we go on to
concentrate on a training or purely
behavioural problem could be negligent and
endanger the animalís life.
problems are generally not abnormal
behaviours. Cats, on the whole, exhibit fewer problems
than dogs, although the evidence is less complete. Up
to one third of all cats and dogs brought to vets for
euthanasia are destroyed because of behaviour problems.
This means that behaviour problems are certainly
the most common cause of death in young animals.
should also bear in mind that pets with behaviour
problems that are tolerated may drastically reduce
the ownerís quality of life.
a number of things to bear in mind when
reading about or discussing behaviour problems with
clients. We are only going to cover these subjects very
briefly. The idea here is to give some idea of the sort of
therapy that can be applied to pets with behaviour problems.
Never jump to conclusions about the nature of a
problem behaviour. Owners often leave important bits of
information out until you have committed yourself to a
diagnosis. Donít give these tips as if they are quick
fixes. Unless you have investigated the case properly
you may well find that the unwelcome behaviours do not
get better but instead get worse. This would be suitably
embarrassing and can even result in legal action.
Sexual problems: Males may lack interest in copulating
with a receptive female. This may be due to several
factors: unfamiliarity with breeding environment, lack of
experience, sometimes a hair ring develops around the
glans penis which prevents intromission, or hormone
levels may be low.
males should be conditioned to breeding
activity by frequently copulating with receptive females.
that have been castrated may masturbate
after being castrated (Schwartz, 1997).
frequent abnormal behaviour in females is
the queen who appears to be in full oestrus but will not
accept the male. Some females show preferences for a
certain male so it is advisable to try the female with a
different stud male (Hart, 1980a).
can experience pseudo-pregnancy if they
ovulate but do not conceive. Robinson (Thorne, 1992)
reports that a good way of reducing the number of
oestrous cycles is to stimulate the vagina with a cotton
bud. Because females may lose condition and even
cause noise pollution with repeated oestrous cycles,
some breeders use this technique.
Excessive grooming: This can cause skin irritation
and loss of hair and is more common in Siamese and
Abyssinian cats. Emotional problems can enhance the
problem and it can be treated with corticosteroids.
the flanks and back may be a displacement
activity displayed when the cat is confused or has
been upset. Excessive grooming may occur if the cat is
continually stressed (e.g., by a new cat in the house).
Self-mutilation, such as grooming to a skin lesion, is
very rare (Bradshaw, 1992).
grooming of her young by an overly anxious
mother results in the newborn being less able
to nurse because they are constantly disturbed, and
they may die due to lack of nourishment and loss of
body heat (Schwartz, 1997).
Prolonged sucking: This can become a vice in adult
pet cats: tactile stimulation of hair and earlobe while
being petted, evoke the nursing response.
kneading with the front paws typically
accompanies sucking (Schwartz, 1997).
Pica: This is defined as an abnormal craving to ingest
unusual substances and is said to occur when inappropriate
objects (such as wool) are ingested (Schwartz,
1997). It is common in Siamese cats. Ingestion of the
material being sucked can result in intestinal obstructions
problem, specifically in cats, can involve a variety
of bizarre substrates, including rubber and electrical
cable but most commonly it involves the sucking and
the ingestion of fabric. It is seen in exotics, especially
Siamese, which suggests a possible genetic link. But it
has also been linked to a traumatic weaning process,
separation anxiety (it can be precipitated by some types
of stress), a deficiency of fibre in the diet and the lack of
any opportunity to perform natural predatory behaviour.
It is a serious problem and can lead to gastric obstructions
and impactions, but also to incredible damage,
especially when the cat generalises from wool to other
for pica is to increase the fibre content of
the diet (sometimes even provide very small quantities
of shredded wool if the cat is looking for lanolin, but also
give occasional laxatives). Restrict access to garments,
and make the fabrics unpalatable, perhaps by sprinkling
with eucalyptus oil.
be a part of a catís normal diet or be
eaten if the cat is nauseous. Grass eating is not a definite
indication of internal parasites, as sick cats may not
eat it and healthy cats may (Schwartz, 1997).
Elimination problems: Elimination in places other than
the litter box is the most common behavioural problem
cited by cat owners (Borchelt and Voith, 1981). These
include urination, defecation and marking problems.
They are usually complex problems which cannot be
cured with any one type of treatment, The client and
veterinarian have to devise strategies
that may include moving the litter box,
placement of food in the area where
spraying occurs thus converting it into a food
area or placing toys in the spraying area to make it into
a play area. Castration may prevent spraying by an
intact male and synthetic progestogens will often suppress
spraying in castrated males or females. Reward
and punishment techniques are not very successful for
Elimination problems may include a refusal to use
dirty litter. Also, physical elimination problems (pain)
may teach cats to avoid the litter (Schwartz, 1997).
If a cat
suffers enough social stress or predisposing
circumstances it may begin to void outside its litter box
multi-cat household, separate litter boxes
should be provided for each cat so that any territorial
conflict can be prevented (Schwartz, 1997).
House-soiling differs from marking in that it involves
the deposition of faeces out of the litter tray and is
usually a product of poor maternal care
and/or poor training. (Persians seem to be
over-represented in this problem.)
ability to discern a litter substrate as being
preferable as a toilet site is a product of the cleanliness
of the nest in which it was raised. Over-fastidious litter
cleaning by the owner can mean that the cat is not easily
able to locate its toilet but, equally, most cats prefer
the litter tray to be well maintained.
relative positions of litter trays and feeding sites
can be crucial in helping the pet to select an appropriate
site. Feed bowls should be in a different room to the
tray. Placing small portions of dried food in areas that
have been fouled tends to deter further mistakes.
Covering the litter box with a dome to make it more private
can help some cats, especially those living in a
household with nosy dogs. Thorough cleaning, as with
marking problems, should discourage return visits to the
Indoor marking: Indoor marking (scratching, spraying
and middening) has to be distinguished in each case
from house soiling. Activities such as claw sharpening
are normal for cats and only become a problem when
the favourite furniture is used. Usually the provision of a
scratching post in place of the furniture solves the problem.
Extreme cases may require declawing. Marking
generally occurs in an attempt to make the perpetrator
feel more secure in its lair and therefore it can include
scratching, urination and open defaecation (i.e., where
no attempt has been made to cover up the faeces, aka
of perceived security in the lair is precipitated
by things such as the introduction of a cat flap, a
young baby, a new spouse or the arrival of new cats in
elimination tend to include areas that either
have a higher concentration of challenging odours, such
as door mats, or those that smell particularly strongly of
the owner, such as chairs and beds, clothes and even
the owners themselves.
and neutered males may urine spray Ė in
other words, the behaviour is not confined to tomcats. It
is just that the behaviour for the females and neutered
males requires a higher threshold than for entire toms.
revolves around making the catís world
more secure, so any form of direct punishment is contraindicated
because it simply makes the cat feel more
If the cat has singled out one new person to be
the target of marking, then it really helps to get that
person to be the only person who feeds
Similarly, encouraging all humans in a household
to use the same soaps, shampoos and scents may help
to homogenise the group and stop one human being
singled out for attention.
Boarding up the cat flap works well but has to
done as part of a campaign that includes thorough
cleaning with a biological detergent followed by surgical
spirit and the placing of dried cat food in favoured sites.
This serves to make the marking points less appealing.
This campaign can be supported with the use of
anxiolytics, tranquilisers Ī pheromonatherapy. These
drugs should be given only transiently and only in
combination with behaviour modification.
Aberrations during parturition and lactation: Some
breeds, like the Siamese, may become very restless
and verge on hysteria. This is not a common occurrence
in other breeds (Joshua, 1968).
first-time mothers may find the whole new
experience overwhelming. If so, the mother should be
separated from the litter to prevent malnutrition or injury
to the young (Schwartz, 1997).
Aggression: This cannot be regarded as abnormal,
but which can be a problem for cat owners. There are
different types of aggressive behaviour in cats (Hart,
Intermale fighting: this can be sometimes eliminated
by castration. Cats are usually extremely flexible
socially, but breakdowns in this ideal
occur when kittens have been deprived of
social learning opportunities between 2
and 7 weeks of age. Intermale rivalry arises and the
intolerance of certain individuals can lead to enormous
problems of aggression.
treatment, these cats can be re-educated by frequent
controlled exposure to the new cats coupled with
distraction techniques, such as feeding, when socialising.
support of the higher ranking individual can
serve to demote the other combatants to the extent that
they are perceived no longer as a worthy opponent by
the alpha cat.
Social-territorial aggression: common aggressive
behaviour, which occurs if a cat, intrudes into another
catís territory. It is often difficult for an adult cat to
accept another cat or kitten into the
household. Synthetic facial gland
pheromones have been shown to have a pacifying
effect on fearful and territorially aggressive cats.
Fear-induced aggression: may be shown when a
child pulls a catís tail or ears or may follow a slap meant
as punishment. Aggression may just be redirected
fright. If made toward another cat, it may initiate
prolonged disagreements (Schwartz,
1997). Do not cut off the retreat of an
aggressive cat or offer direct challenges,
such as staring.
types of aggression may be controlled by
drugs but others may only be controlled by changing the
situation, e.g., an established adult cat directing
aggression towards a new cat.
multi-cat household the same cat is always the
first victim of aggressive outbursts by other cats. If this
happens, the cats should be separated as soon as possible
after the first attack to prevent further occurrences
Aggression towards visitors and owners: this can be
a result of insufficient socialisation and gentling,
causing a display of defence aggression
or predatory chasing, for example, of
human feet. Or it can be redirected
aggression towards humans by cats that have just seen
conspecific rivals through a window.
Treatment is for the owner to make time for some
structured play with the cat and, perhaps, consider
introducing a less threatening cat, such as a kitten of the
opposite sex, into the home.
Overeating: This is often associated with overfeeding
by the owners, especially when they have assumed that
vocalisation by the cat is a sign of hunger.
Over-attachment: If this is the problem, the cat
may follow the owner constantly, be upset
in his/her absence and have a tendency to
suck skin and clothing.
for over-attached cats is to keep petting
to a short duration and only at the initiation of the owner;
in other words, stop rewarding attention-seeking behaviour
with petting. Many of these cats quickly learn just
how to get attention on demand. Periods of isolation in
an appealing room should be increased from a couple
of minutes to six hours over a period of two or three
months. In other words, a process of habituation.
and separation anxiety: These can
result from a lack of early exposure to
appropriate stimuli during socialisation periods, a traumatic
incident, or old age. Cats in this class often
appear withdrawn, have a low crouching gait and are
reluctant to enter open spaces. They may also show
psychogenic vomiting and diarrhoea, and may overgroom
themselves or self-mutilate.
specific stimulus is responsible for the behaviour,
then systematic desensitisation is indicated. Where
there is generalised nervousness, drug support is
sometimes helpful. Specifically, over-grooming and
selfmutilation have their origins in
stress. When a cat comes up against a
confusing or frustrating situation, it
will often stop whatever itís doing and groom itself. The
stress of isolation for instance can send cats into a
grooming obsession in the same way that a flea allergy
can. Ultimately, the danger is that trauma to the skin
ends up generating endorphins that mediate a form of
self-reward to the cat and the cycle tends to perpetuate
itself. Treatment must centre on reducing the stressors
in the catís life and, possibly, the use of drug support
such as benzodiazepines (valium) or tricyclic antidepressants
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