Friday, April 29, 2011

The Mysterious Behavior of the Cat

The cat is a much loved and well-known animal. In Western countries it has become one of the most popular companions; on farms it has been valued as a rodent catcher for centuries. Loved and familiar though it is, the cat remains an enigma. He is friendly to people and yet, Rudyard Kipling said he "walks by himself", accepting comforts of the human home while behaving as though he is fully independent. These paradoxical qualities cause some to mistrust and even hate cats. The cat, more than any other domestic animal, has been as much persecuted as it has been appreciated. It is featured in fables and myths. Even many cat lovers treat them as mysterious. However, in a time in which a great deal has been discovered about the biology of behavior, many of the cat's former secrets have been revealed. Cat people often make careful observations of their cats, but most people also appreciate that each cat has a distinct personality. It is hard and often misleading to make generalizations about the cat. The body of knowledge has expanded in recent years, so that more statements  backed with evidence can now be made both about the common features of domestic cats and about the origins of their differences. Interest in the cat is widespread due to its predatory activities and abilities and its popularity as a companion animal.

Though a lot has been learned in the last few years about the behavioral biology of the cat, a great deal more remains to be discovered. Whether or not the cat walks by himself, he still keeps some of his secrets to himself.

As a cat grows up, its behavior develops with regularity and consistency. Cats are adaptable and changeable in their behavior, responding sensitively to changes in their environment. They are highly variable in their habits. Some cats spend much of their time hunting, while others rarely leave the comfort of the armchair. Their behavior depends on factors that are inherited, and on the cat's experience, a part of which it actively seeks.  Early handling has a number of effects on the behavior and physical development of cats; handled animals tend to develop more quickly. The quality of early nutrition has effects on development as well.

Cat Relationships
Cats form strong social relationships with familiar individuals, usually close kin. In groups of feral cats, kittens often suckle from females other than their mother. Humans and members of other species can be included in the social group and responded to with affection if they are encountered by the cat when it is young. Despite a basic ability to respond socially towards people, adult cats and kittens show individual variation in their friendliness towards humans, whether familiar or unfamiliar.

Kittens who are separated from their mother too early (at least 2 weeks) develop a variety of behavioral, emotional and physical abnormalities. They become unusually fearful and aggressive toward other cats and people, show large amounts of random and undirected locomotor activity, and do not learn as well. Some develop asthma-like respiratory disorders.

Kittens raised with other species, ie rats, never kill rats of the same strain when they grow up. Kittens whose social companions during early life were rats form social attachments to rats, inhibiting later predatory responses to them. However, when given the opportunity to form social bonds with other kittens as well as rats, other kittens are preferred.  They show a distinct tolerance of rats and reduced predatory response towards them. Willingness to try new food and preferences for specific types of food appear to be strongly influenced by the mother. When given a new food with the mother, they eat it. When on their own, they do not eat it until they have been exposed to it for a longer time. Young cats are well adapted to learning from their mother, and show a strong interest in and ability to learn from the behavior of other cats. This is generally found in many species and is known as social learning. Kittens usually kill the type of rat they have seen their mother kill when young.

Social Learning
Adult cats also show social learning. Kittens who watch their mother perform a task acquire the response quickly, but kittens who acquire the response by trial and error do not. Kittens watching the mother learn sooner than kittens who observe a strange female; social leaning works if the model cat is familiar to the observer.

Cats observe the actions of a human leaving the room, but acquire some learned responses faster by observing another cat than by conventional conditioning practices. Observing another cat acquire the response important, and has a more beneficial effect than watching another cat perform a skilled response that has already been learned.

Siblings are important in social development. Social play and object play develop at different times, showing that these forms of play are separately organized and controlled. Even in terms of the motor patterns used, object and social play differ distinctly in a number of ways, e.g. repetition of certain motor patterns occurs frequently during object play but rarely during social play.
Play is not needed for at least the basic element of predatory behaviors to develop. Play may have subtle beneficial effects on predatory skills.

Behavioral Traits
The kitten's development is well organized. The systems that generate the intricately integrated behavior of an adult cat seem to be goal-directed and are resilient to both internal and external disturbances. Most cats eventually become reasonably competent predators, regardless of the experience they have as kittens. Cats are versatile and adaptable, able to thrive in different environments.

Cats are individuals with distinct characters, similar to other cats in general responses, but different in others. Many cat people perceive their cat as a unique individual, as reflected in the popular literature on the cat, in which references to individuality and personality are common. There may be a link between coat color and certain behavioral characteristics.

In human personality, most behavioral traits can be grouped in to five broad categories known as "the Big Five" personality factors: Extraversion, Agreeableness, Conscientiousness, Neuroticism, Openness to Experiences. Maybe more relevant to studies of animals, child psychologists have identified a limited number of basic elements of personality: emotionality, sociability and activity. The categories are activity, reactivity and avoidance in octopi and confident, excited and sociable in rhesus monkeys. There are active and passive coping strategies, clusters of behavior shown in response to challenging situations, and the extent to which there are found as distinct response styles with a specific stimulus.

Genetic Factors

Genetic factors may exert some influence over behavior, which contributes to individuality of the cat. There are also behavioral difference between various cat breeds. Cat breeds are genetically distinct at certain loci, the differences in aspects of their behavioral style indicate that genetic factors may underlie some of the observed individual variation in behavior.

Another line of evidence linking variation in behavior to genetic variation comes from observations of behavioral difference between cats with different coat coloring. For example, cats carrying the non-agouti allele usually black, may be more tolerant of crowding and the conditions of urban life than those carrying the agouti allele. Red, cream or tortoiseshell colored kittens struggle for a longer time and make more escape attempts when handled by an unfamiliar person than kittens with other coat colors .
This line of thinking was extended by linking coat color polymorphisms and their underlying genetic profiles, to behavioral differences and their subsequent impact on social structure. They suggest that the orange allele may be linked to aggression in males, while some other mutations, such as the non-agouti, are linked to greater amicability and grouping tendencies. This explains why the orange alleles tend to be found at relatively low frequencies, at least in urban environments, compared with other more recent mutations, such as the non-agouti and the blotched tabby.  In urban environments, where cat densities can be very high, aggression is not related to reproductive success of males as a result of the promiscuous mating system. Aggressive males can actually lose mating opportunities due to their intolerance of other male cats and because they spend their time fighting rather than mating. In contrast, successful males mat be seen sitting patiently waiting for their opportunity to mate a female in oestrous, and sperm competition, rather than physical combativeness, seems to be the main mechanism in male mating success. Thus, aggressive males with the orange allele will on average have a relatively low degree of mating success. Despite being one of the most ancient coat color mutations, the orange allele has a small frequency.

In contast. rural cat populations tend to be at lower densities, with cats more widely spread. Dominant males can monopolise females more, and aggression may be more successful. Rather than a promiscuous mating system, a relatively high degree of polygyny appears to operate in rural areas. The orange allele is more common in rural environments. In certain circumstances, genetically linked behavioral traits may be favored.

A relationship between coat coloring and behavior appears to occur in a variety of other species, and a number of possible reasons for their occurrence have been suggested. Changes in coloring may directly influence the function of sensory organs. i.e. lack of protective pigment in the iris of albino animals leads to vision problems, especially in bright light, which may underlie some of their common behavioral traits, such as sluggishness of reaction. The mechanism underling coat color is linked to those underlying the control of behavior. Coat coloring pigments (melanins) are produced by the same biochemical pathways as the catecholamines, such as dopamine, which are important in brain activity.

Genes controlling coat colour are located at positions on chromosomes close to other genes which have some influence on the function of the nervous system,  i.e. blue-eyed white cats and white cats with orange eyes are often deaf. Genetically-induced defects affect behavioral. The gene involved in the production of the white coat color my be positioned close to a gene which induces deafness in one or both ears, thus cats inherit both traits.

A kitten's early environment has a strong influence on its behavior later in life. Behavioral measures which are indicative of the characteristics of boldness and nervousness are influenced by a variety of different types of early experience. Kittens handled regularly during the first 45 days of life approach unfamiliar objects more rapidly, and spend more time close to them than non handled animals. Variation in boldness or nervousness may be directly related to variation in forms of early experience including the social environment and exposure to handling by humans, These aspects of early experience also have effects on other types of behavior.  Human handling (socialization) timing, amount of early handling of a kitten and the number of handlers influence its later friendliness to humans.

It may affect how kittens adapt to housing in animal shelters, cats who are not socialized with other cats show more signs of distress when group housed in a shelter than socialised cats, cats not socialised to people are more stressed in single and group housing than socialised cats. Cats can be described as tense, high strung, calm or anything else. There is a huge variation. Many different types of early experience can exert some influence on the expression of a specific behavior pattern at a later date; one aspect of early experience can influence a number of different types of behavioral characteristics.

Paternity and Friendliness
Genetic and environmental factors interact to influence the development of numerous behaviors in the cat. One factor which causes variation in kitten friendliness is paternity. Kittens of different fathers show varied friendliness, though they never see the father. Genetic factors caused this. A friendly father produces kittens who are more friendly, than does the less friendly father. The father’s friendliness when young and not when an adult can more accurately predict the friendliness of his kittens. Genes do not code for behavior patterns. Genes from the father generate a difference in behavioral characteristics by indirect routes, i.e. by affecting growth rate, which may in turn affect socialisation to humans in a colony situation and affect subsequent friendliness.
Variation in kitten friendliness is partly explained by kitten paternity. Paternal effects on kitten behavior may be attributed to breed differences between fathers, also, variation in early handling affects subsequent behaviors. Friendly father’s offspring who are handled during early life can be quicker to approach a person and spend more time with them than non handled kittens and the offspring of an unfriendly father. A friendly father’s offspring are quicker to approach and explore new objects. The effect on friendliness may be a more general effect on boldness, due to paternity, influencing a general response to unfamiliar or novel stimuli, regardless of whether it is a person or inanimate object.  Early handling has a specific effect on the cat's behavior towards humans-its friendliness. There is a degree of additive effects in the genetic and environmental effects on some of the cat's responses to humans. Friendly-fathered, handled cats behave in the most positive way to people, from the interactive effects of genetic and environmental factors on individual behavioral characteristics. The paternal effect on friendliness or boldness may only become expressed at a certain point of development and maturity.

Differences Along the Timeline
Early individual differences in predatoriness do not necessarily predict subsequent variation in the predatory abilities of older cats.  Variability of individual characteristics early in life, around 3 months, may not always predict subsequent individuality in the cat.

There are a number of other reasons why individual behavioral characteristics at one time may not relate to those at another, i.e. cats may develop at different rates and these differences may be clear at one age but not at another. The behavioral expression of particular characteristics may change with age, as may the factors controlling and functions of particular behaviors. In contrast, some studies have shown cross-time consistency in cat individual characteristics. Boldness and curiosity remain reasonably stable despite changing social environments. When cats are tested in groups rather than singly, they behave differently, showing that individual differences in behavior in a non-social context may become amplified when individuals are allowed to interact with each other. There develops a clear social ranking, which may influence individual behavior and cause the consistency of the patterns within a group. Friendliness is attributed to quick/noisy kittens who are more exploratory and quickest to investigate unfamiliar stimuli. These differences can be present from birth and persist into adulthood. Characteristics are also open to environmental influences.

Individual variation in characteristics, such as timidity and boldness, are present at birth or appear shortly after birth. It has been shown in different species that they are related to constitutional differences in the physiological characteristics of the individuals.  In wolves, the dominant cub in a litter differs in heart rate and other autonomic process in some test situations. Physiological systems change with time and situation as do behavioral ones.  Physiological differences are related to individual differences in behavior. There may be a physiological basis for certain personality traits, especially those present at birth and behavioral stability across time could be attributed to stability of physiological systems.  Certain measures of behavior change dramatically according to the contexts in which they are observed, while others are less variable.

Special Circumstances
When midway through a quarantine, cats are less relaxed, playful and more aggressive and nervous than prior. At release, they are more friendly, affectionate and timid. Three months later, they are more affectionate, nervous and vocal than before quarantine.  The interactions that their people could have with their cats were much more restricted in the small quarantine cages than at home. The experiences changed their relationship with their cat; they may have had expectations of how quarantine would affect the cats, and may have changed their behavior and perceptions accordingly. Time and context changes during the quarantine. Some of the changes may reflect true changes in cat individuality. Important real life contextual changes, such as moving and being quarantined or housed in a cattery may result in change in a cat's behavioral style. The context can dramatically affect a cat’s individual character. An aloof, placid and low-ranking cat can become aggressive, energetic and dominant if two cats must compete for one food reward. This has a marked effect on behavior, seen in social competition, even in the same individuals.

Relationship Between Cats and People
Relationships between cats and people may be especially open to contextual influence because in a social situation, the behavior of others has a strong effect on an individual's own behavior. Individuals may also develop complicated context-specific ways of behaving according to past experience with other individuals which act to conceal or constrain the expression of any underlying stable behavioral characteristics.  Contextual effects on individuality can be external contact or internal (e.g. hormonal) state of the animal.

Behavior Types
These terms are used to describe the behavior in a number of species, to describe individual differences: active/reactive, social and calm, which may underlie many of the behavioral differences in between individuals.  Others are active/aggressive, timid/nervous and confident/easy going, though some individuals do not fit into these groupings, showing that there is unlikely to be a limited number of easily definable cat personality types. An expression of friendliness could be the result of an interaction between an individual's general boldness in novel or challenging situations, its specific attraction to humans, both of which are affected by a variety of factors, and influenced by the current situation. There is a complex origin and development of individuality. Situational stability and different behavioral styles help cats to cope with caging or quarantine. This shows a new approach to understanding the consequences of these situations, which has important implications for animal welfare.

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Singing Nettles Herbal Clinic by Amanda Dainow is licensed under a Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License.
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