Cat Behavior PDF
Cat Behavior PDF
Cat Behavior PDF
DOI 10.1007/s10071-015-0897-6
REVIEW
Introduction
At first glance, it may seem that any member of the family
Felidae would make an unlikely human companion. Most
members of Felidae lead solitary lives and only engage in
social behaviors during mating or kitten rearing (Macdonald et al. 2000). The only members of Felidae that
commonly live in social groups are lions (Panthera leo),
cheetahs (Acinonyx jubatus), and the facultatively social
domestic cat (Felis silvestris catus), which can display
varying levels of non-obligatory social behavior depending
on environment and upbringing (Leyhausen 1988; Mellen
1993; Bradshaw and Cameron-Beaumont 2000; Turner
2014). So, how did the domestic cat become one of the
worlds most popular pets, with over 600 million cats living among humans worldwide (Driscoll et al. 2009)?
We may have a long way to go before we know. While
research on domestic cat behavior and cognition is growing, many questions remain unanswered. How the development of cat behavior and cognition is influenced by
factors such as species-specific biological predispositions,
domestication and lifetime experiences, including the
humancat bond, remains largely unexplored. In general,
comparatively little research has been devoted to cat cognition, especially when compared to our other popular
companion, the domestic dog (Udell et al. 2010; Merola
et al. 2015). Nonetheless, research on cat cognition could
have important theoretical value and management and
welfare implications, including improved humancat
interactions (Bernstein 2014). Therefore, the purpose of
this review is to survey the current status of several areas of
cat cognition research, broadly defined as all ways in
which animals take in information through the senses,
process, retain and decide to act on it (see Shettleworth
2001, p. 277), that have received at least some scientific
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Perception
Cat sensation and perception have historically received
more attention than other areas of cat cognition. The
majority of this research has examined cat auditory perception (Gerken and Sandlin 1977; Costalupes 1983;
Heffner and Heffner 1985; Rauschecker and Kniepert
1994; Cornwell et al. 1998; Malhotra et al. 2004; Witte and
Kipke 2005), olfactory perception (Willey 1973; Verberne
and de Boer 1976; Verberne 1976; Won et al. 1997;
Mermet et al. 2007), visual perception (Blakemore and
Cooper 1970, 1971; Peck and Blakemore 1975; Blakemore
and Van Sluyters 1975; Blake and Hirsch 1975; Buisseret
and Singer 1983; Cornwell et al. 1998), and cutaneous
sensory mechanisms, including vibrissae, or whiskers,
which aid in movement in low-light conditions (Bradshaw
et al. 2012) and supplement for the cats lack of shortdistance vision (Williams and Kramer 2010).
Research on cat perception has demonstrated how early
sensory experiences influence brain development and perception (Blakemore and Cooper 1970; Blakemore and Van
Sluyters 1975; Blake and Hirsch 1975; Korte and Rauschecker 1993; Burnat et al. 2002, 2005). One such classic
experiment on vision by Blakemore and Cooper (1970)
found kittens selectively reared in an apparatus covered with
either vertical or horizontal lines could not detect the
opposite line orientation to the one in which they were reared
under, providing evidence for environment-dependent neural plasticity. Additionally, Blake and Hirsch (1975) found
that monocularly deprived kittens had a reduced proportion
of binocular cortical cells and impaired binocular depth
perception, providing evidence that early experience is
critical in developing cells used for binocular depth and
stereopsis. Rearing environment also influences perception.
Laboratory-reared cats have demonstrated an impairment in
response and discrimination learning to visual stimuli
_
(Zernicki
1993), an impairment of visual associative learn_
ing (Zernicki 1999), and delayed response to auditory cues
(Wikmark and Warren 1972) as compared to free-roaming
cats, indicating that poor rearing environment can influence
perceptual ability later in life. Training has also been found
to improve performance on visual perceptual tasks (Sasaki
et al. 2010), such as an increased contrast sensitivity to
specific frequencies of gratings accompanied by changes in
the primary visual cortex (Hua et al. 2010).
While much of this research originated from scientific
interest in the development of sensory systems and the
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Physical causality
The ability to understand cause and effect is of great
adaptive advantage as the use of causal principles allows for
the transfer of learning gained in one situation to a novel
situation (Seed et al. 2006). Whitt et al. (2009) studied the
extent to which cats understand physical causality through
use of the string-pulling test. Cats were first trained to
obtain a piece of food attached to a short string placed
perpendicularly under a box, with only the tip of the string
visible to the cat. They were then presented with different
arrangements of strings including one long string, two
parallel strings and two crossed strings. In arrangements
with two strings, food was attached to only one string. Cats
had ten attempts to retrieve the food successfully by pulling
on the correct string. Although cats successfully pulled the
single string to receive the food, they were unable to perform above chance when presented with either of the twostring arrangements. This finding suggests that the cats did
not understand the function of the string in its means-end
connection with the food reward. However, as this area of
cognition has received sparse attention, more research must
be conducted to fully examine the abilities of cats in this
domain, especially to ensure confirmation or development
of species appropriate methodology. For example, Whitt
et al. (2009) suggested that the cats may have found stringpulling rewarding in itself, whether or not they received the
food. As Whitt et al. (2009, p. 742) state, It is therefore
important to investigate cognitive abilities based on their
importance for the differing ecological needs of species.
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Social cognition
Many domestic cats live socially and rely on communication within and between species to thrive in a variety of
environments; however, science is only in the beginning
stages of understanding cat sociality and social communication (Bradshaw et al. 2012). In domestic cats, which
display plasticity in whether or not they live socially, the
formation of social groups is dependent on the availability
of food, shelter and mates (Liberg et al. 2000; Macdonald
et al. 2000; Turner 2014). Although many think of cats as
solitary, research indicates free-roaming domestic cat
social groups are not random aggregations around a food
source (Macdonald et al. 2000; Denny et al. 2002), rather
cats come into proximity and engage in social behaviors
with preferred associates non-randomly (Curtis et al. 2003;
see also unpublished thesis work by Wolfe 2001 and
Shreve 2014). Additionally, cats display a wide array of
affiliative, aggressive and investigatory behaviors toward
conspecifics (Bradshaw and Cameron-Beaumont 2000;
Curtis et al. 2003). Likewise, cats enter into a range of
relationships with humans. Some live as companion animals in human homes, and others live as strays which
navigate our urban spaces, often dependent on human
caretakers for resources. Pet cats have countless social
encounters with humans (as well as other household
members), with varying levels of complexity and success,
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Attachment
Attachment is an enduring affiliative social bond formed
between an animal and a specific individual (Ainsworth
and Bell 1970). In attachment, the animal behaves in ways
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Cat personality
Researchers across numerous disciplines have begun to
explore individual differences in animal behavior, using
various terms such as personality, individuality, temperament, distinctiveness, behavioral style or behavioral syndrome, (Gosling 2001, 2008; Raihani et al. 2014). While
researchers have not always agreed on the correct terminology to describe the occurrence of individual differences
within non-human animals, for consistency we will use the
definitions provided by Gosling (2001) when referring to
personality and temperament throughout. According to
Gosling (2001) personality can be defined as those characteristics of individuals that describe and account for
consistent patterns of feeling, thinking and behaving (p.
46); in other words, a prolonged state in which behavioral
patterns are relatively consistent over time and various
circumstances but can be influenced within the life of the
animal. A closely associated idea is that of temperament,
which Gosling describes as inherited, early appearing
tendencies that continue throughout life and serve as the
foundation for personality (p. 46). In this case, temperament can be used to describe the biological dispositions of
the animal (Wynne and Udell 2013).
Cat owners and even researchers working with cats
often indicate that they feel cats have definite personalities.
For example, Feaver et al. (1986, p. 1016) stated we felt
that each animal in our laboratory colony had a distinct
personality in the sense that the sum total of its behaviour
gave it an identifiable style. However, is this feeling
supported by empirical science? Durr and Smith (1997)
examined whether stable individual differences in cat
temperament (a measure of personality as defined by
Gosling) could be identified in testing situations. Study cats
were first observed in their normal living conditions to
provide an initial assessment of their behavior, including
order of approach, latency, and attention span. Cats then
experienced various conditions, including disruption of
their social environment via confinement or overcrowding,
to determine whether a stable environment is important in
maintaining individual differences in cats. In additional
tests, cats were presented with novel stimuli, including a
moving stimulus that emitted moderate to loud noises (e.g.,
remote-controlled car with a motor), an unfamiliar animal
(e.g., dog, rabbit), and an intense novel stimulus (e.g.,
vacuum cleaner that was turned on). Cats were also
deprived of food and then presented with a highly desirable
food item. Durr and Smith (1997) found that the behavioral
responses of cats were consistent despite changes in their
environment, indicating the stability of the social environment is not crucial in maintaining the stability of individual behavioral responses. The authors concluded
individuality is the expression of inherent temperament
and not simply the release of responses created by a certain
set of stable environmental variables (Durr and Smith
1997, p. 416).
Other studies have also found evidence of stable personalities in cats. For example, Turner et al. (1986) found
that the behavioral trait of friendliness was consistent in
kittens aged 38 months. Lowe and Bradshaw (2001)
found that four distinct personality styles were stable
between 4 and 24 months of age (discussed further below).
Additionally, Durr and Smith (1997) found that cat group
structure is reliant on the individual characteristics of the
cats in the group, indicating that individual differences
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between cats can be influenced by conspecific social relationships and potential human relationships as well.
Studies have determined personality styles of cats utilizing various methodologies, such as human ranking of cat
differences through direct observation of their behavior
(Feaver et al. 1986; Turner et al. 1986; Raihani et al. 2014)
or exposure to various conditions (novel stimuli, unfamiliar
persons, increased socialization) with the cats behavioral
responses to these conditions recorded (Meier and Turner
1985; Mertens and Turner 1988; McCune 1995; Durr and
Smith 1997). Combined, these studies have described many
personality types within cats. Two traits commonly found
include boldness, which describe an individual who has
actively approached novel stimuli in the past and would be
expected to do the same in the future, or shyness, which
may describe an individual who has reluctantly or not
approached novel stimuli in the past and would be expected
to do the same in the future (Mendl and Harcourt 2000;
Gosling 2001). Bradshaw et al. (2012) divided cat personality into three types, based on findings from previous
research. The first personality type involves an individual
with traits such as sociable, confident, easy-going,
trusting, bold and initiates friendly interactions
(Meier and Turner 1985; Feaver et al. 1986; Mertens and
Turner 1988; McCune 1995). The second personality type
involves an individual with traits such as timid, nervous,
shy and unfriendly (Meier and Turner 1985; Feaver et al.
1986; Mertens and Turner 1988), and the final personality
type involves an individual with aggressive traits (Feaver
et al. 1986). In addition, Lowe and Bradshaw (2001)
described four distinct behavioral styles (a term they use
interchangeably with personality) that exist in cats. These
include the tendency to stay indoors, to rub (people or
objects), to investigate and the tendency to boldness. But,
other personality styles have also been described in cats
(Mendl and Harcourt 1988, 2000; Bradshaw et al. 2012).
More research may be needed to establish a consensus
about which terminology, personality styles or traits have
the most predictive value or whether some of these categories might be combined to establish a concise consistent
categorization of feline behavior types.
Researchers have also questioned the point in development when differences in cat personalities arise. Raihani
et al. (2014) found there were intra-litter differences in
kitten behavior at as early as 56 days old, and kittens at
34 weeks old already display relatively stable behavioral
differences. This indicates that differences in personality
existing early in development maybe even from birth
although this study did not examine kitten behavior prior to
day 5, as to not disturb the early motherkitten relationship.
The researchers also noted they did not initially expect to
find individual differences at such an early age. This
finding may suggest there is a genetic component to
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Conclusion
We still have a long way to go until we have an inclusive
body of research on cat cognition. Many questions are still
largely unexplored. To what extent do cats alter their social
behaviors for communication with humans? Are there
cognitive differences between various groups of cats (feral,
shelter, pet)? How do lifetime experiences, such as training, influence cat cognition? Exploration of this research
will contribute to our scientific understanding of how
domestication, the human bond and adaptation from a
solitary to social lifestyle influence cognition. This research
could also have important applied applications; for example, free-roaming cat management is a topic of debate that
may benefit from a greater understanding of how cats
navigate and perceive human spaces. Additionally, cat
cognition research may lead to improved humancat
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