School of Clinical Veterinary Science, University of Bristol, Langford, Bristol, UK
There are many reasons why vets should think about the complex behavioural life of the domestic cat. These include:
A responsibility for welfare. As vets we tend to concentrate on the physical welfare of our patients, but their psychological well-being has an equal impact on their quality of life. Cats which live in unsuitable or stressful environments, or which cannot show their normal patterns of behaviour can suffer prolonged periods of poor welfare.
Giving preventative advice. Vets are in the unique position of seeing many owners soon after they acquire a kitten at first or second vaccination. The first weeks of life are a very important period in the behavioural development of kittens, and giving owners the right information at this time can impact on the cat's welfare, and the bond formed between cat and owner, for the rest of the animal's life.
Identifying where stress may be important in disease states. It is now well known that stress occurring in response to environmental factors interacts with physical health. Acute stress has an impact on susceptibility to disease, for example. Stress also has an influence on recovering and rehabilitation after disease, injury or surgery.
Handling cats in the practice. The welfare of cats in the veterinary practice is influenced by an understanding of their behavioural needs.
Behaviour 'problems' in cats are common, and owners frequently ask for help with them from their vet.
The first aspect of understanding cats is to consider their ethology, or natural patterns of behaviour. Despite being a common domestic pet, relatively little research has focused on 'normal' cat behaviour, and there remain debates and controversies about various aspects of their behavioural repertoire. This presentation highlights some of the important characteristics of the behaviour of the domestic cat.
Domestication of the cat was relatively recent (approx. 6–8,000 years ago) compared to that of the dog. Furthermore, reproductive activity in cats was not controlled in domestic cats to the extent that it has been in dogs. This means that the modern day domestic short haired cat shares many behavioural characteristics with its ancestral species, the African Wild Cat. African Wild Cats still survive in some areas of Africa, and we can gain some insights into the factors influencing the behaviour of our modern cats by examining their behaviour. Because they hunt alone, and have prey species dispersed over a wide area, African Wild Cats spend a high proportion of their time engaged in prey seeking or hunting activity. Because hunting is often unsuccessful, this activity is maintained even after a successful kill and when the cat has eaten - hence satiety does not reduce motivation to hunt in the same way as it does in canids. The African Wild Cat is also a largely solitary species. They do not hunt co-operatively and usually only live in groups where a sufficient supply of food is available within a concentrated area. Whilst related female cats may form social "colonies" under conditions of plentiful food supply, and show a range of friendly behaviours within this context, they remain antagonistic to individuals from neighbouring colonies.
The behaviour of the domestic cat is also similar to their wild ancestors in other respects, including the establishment and maintenance of territories, the size of which varies according to the availability of resources and presence/absence of other cats. Home ranges consist of a core area where the cat will usually be able to relax, and do activities such as sleeping and playing. Prey items are also usually brought back to the core area to be consumed. The range will also usually have a wider area that the cat occupies, for example, when hunting. Particularly in domestic situations, this space is often shared with other cats, often using a 'time share' system to avoid face-to-face meetings.
The other important characteristics of African Wild Cats which have a profound influence on the behaviour of modern domestic cats relates to their senses and communication strategies. For example, the visual abilities of cats are adapted for predation in poor light conditions: they have excellent light gathering power and movement detection, but relatively poor colour vision and a slow accommodation for focus. Scent is a very important sense for cats, and gives them a whole world of information that we are completely unaware of. As well as an acute sense of smell, cats use the vomero-nasal organ in the roof of the mouth to 'test' chemical signals in the air, with the ability to pick up scents that are only a few parts in a million.
Although cats do show obvious visual signalling for communication, this is generally less subtle than that shown by species that have evolved as social predators, like humans and dogs. Signals in cats tend to be 'obvious' indicators of defensive threat (e.g., fluffing up and showing the teeth), or friendly approach (e.g., the 'tail up' signal). This is because there has been no evolutionary pressure to develop the kinds of complex visual signalling that social predators need to convey complex information between group members for successful hunting and group cohesion.
Olfactory signals are very important for cats both for communication and orientation. Cats have scent glands all over their body, and signals are deposited both on physical aspects of their environment and social contacts (cats, humans and other species). The social function of scent signals is to recognise members of the same social group, but also to maintain distance between individuals that are not group members. Scent signals also provide information for the cat itself: leaving signals that have different 'meanings' for them around their environment gives them some control over their environment. For example facial rubbing is used in areas of the territory where the cat feels comfortable and unthreatened (i.e., the core area), whereas signals such as urine spraying are indicators of potential threat. Urine spray marks are generally left in response to some kind of change in the environment, particularly related to fluctuations in local social dynamics between cats. Because urine spray marks provide a valuable indicator to the cats of where to be alert, it is adaptive for them to 'top up' scent marks as they fade.
Another aspect of cat behaviour linked to their ancestry is their toileting behaviour. Although inappropriate toileting is a common 'problem' for cat owners, changes in toileting site occur for good 'cat reasons' for the cats themselves. Once established, toileting locations and substrates do not change if nothing in the environment changes - changes will only occur where either location or substrate are no longer suitable for the cat.
Cats' criteria for a suitable toileting location include:
A quiet secluded area
Suitable material for burying
Substrate preference often acquired as kittens
Separate locations for urine and faeces
Although many of the characteristics of cats are maintained from their ancestral species, there are some aspects that have changed over the process of domestication. The most significant of these are patterns of social behaviour and groupings in cats. Wild cats were likely to be initially attracted to human settlements because of the increased availability and density of rodents around grain stores. However, in order to take advantage of this resource, cats had to be more tolerant of close proximity to both people and other cats. With factors such as increased isolation associated with feeding in temples and religious status, these cats become progressively more isolated from the 'wild' population, resulting in differences in social tolerance. Understanding the behaviour of modern feral cat colonies helps unravel some of the complexities in social living that impact on cats living in domestic environments. Feral group size depends on food availability, and groups are generally made up of related individuals. These groups show many friendly behaviours with aggression between group members being rare. However, group cohesion is maintained by members knowing each other well and being able to predict behaviour patterns and responses, plus a limited need for any conflict over resources. These limitations mean that there can be problems in multi-cat households where individuals are introduced as adults and/or where there is competition for space or other important resources. Problems between cats, such as aggression or other behavioural signs of stress, occur most often where individuals do not regard each other as part of the same social group. Identifying differences in social compatibility is therefore key to ensuring the welfare of domestic cats kept together in groups.
References
1. Bradshaw JWS, Casey RA, Brown SL. The Behaviour of the Domestic Cat. 2nd ed. Wallingford UK: CABI; 2012.