Description of the Feeding Behavior of the Cat

2. Description of the Feeding Behavior of the Cat

Predation and Hunting

Unlike the domestic dog, the body type of the domestic cat is not far from its wild ancestors. However, differences in prey sizes have led to significant differences, e.g: domestic cats use their incisors less, meals are more frequent and their way of consuming prey is also different: domestic cats begin with the head, large felids begin with the viscera.

Hunting Instinct or Learned Behavior?

Predatory behavior is innate: all cats probably know how to hunt, but certain aspects seem to be learned. Approach and pursuit are stimulated by littermates. Hunting behavior is more likely seen in kittens from a queen that hunts. Kittens learn to catch and to kill the same prey that their mother hunts (Bateson & Bateson, 2002).

During weaning, there is an amazing training program for hunting displayed by the queen:
- 4th week: the queen brings meat pieces to the kittens
- 5th week: she eats dead prey in front of her kittens
- 6th or 7th week: she lets them eat the prey
- 8th week: she brings a live prey in order for the young hunters to learn to kill.

The first hunting sessions occur at 3 months. At 4 months of age, the young hunters are confirmed. The absence of predatory experience does not seem to interfere with motor abilities but often reveals prey selectivity issues. A kitten must indeed be taught that a mouse can be eaten. If it is not done before the age of 3 months old, the cat can starve even in the presence of the unknown prey! However, even cats that did not have access to prey when they were kittens can learn to become proficient hunters.

It has been speculated that feeding a cat may reduce its desire to hunt, but evidence to support this is lacking or controversial. Cats that are provisioned with food at home spend less time hunting than cats that are not provisioned with food, but both still hunt even if the number of actual prey caught and consumed is hard to quantify (Fitzgerald & Turner, 2000).

Hunting Session

Unlike dogs who hunt in packs, cats in the wild are solitary and opportunistic hunters. They catch small prey and eat alone. Observations show that they often fail in their attempts to catch prey: only 13% of tracked prey is actually caught (Kays & DeWan, 2004), every success representing 3 - 5 attempts (Fitzgerald & Turner, 2000). A cat brings home an average 0.7 prey a week (Woods et al, 2003).

Hunting sessions can last 30 minutes, on distances between 600 and 1800 m in their territory. An obvious variability exists between individuals: for example, male cats hunt longer and further than female cats.

Cats spend two-thirds of their awake time to hunt in natural conditions. Hunting behavior is composed of several sequences:
- Stalking the prey
- Approaching and pursuing
- Catching the prey by a central leap (their body will be low to the ground and they move slowly toward the prey, pausing prior to leaping to attack)
- Killing by biting the neck, following an eventual fight
- Consumption rarely occurs at the location of the catch (for the reason of quietness).

The domestic cat is a member of the Felidae family, Felis catus and is a strict carnivore. (©J.-P. Lenafant/RC/Norwegian Forest Cat).

As the process progresses, the sequences are less and less modifiable by the cat’s experience or it’s’ environment. The first steps (seeking, stalking and approach) are indeed flexible as a result of adaptation to different situations, while the last ones (e.g. attack and bite) are more stereotyped to secure efficient catch and kill, and thus individual survival.

Cats rarely bury their catch for postponed consumption. Cats eat rather quickly and then regurgitate furs and bones. Prey cleaning is rather poor, unless it is voluminous (such as pigeons or young rabbits). The cat breaks the bones and masticates with its large premolars. It can eat an entire mouse in less than one minute. When consuming a mouse, it starts at the head and eats in the direction of the tail (Case, 2003). It is often the logical consequence of the killing method, during which the neck is broken. It might also be related to an adaptative behavior, securing prey catch and intake.

Leyhausen (1979) (quoted by Rabot, 1994) has shown that the complete hunting sequence is in fact controlled by a system of progressive and different motivation phases:

- Interest is awaken by auditory stimuli (scratching, grating), which enables the cat to locate precisely the prey. In veterinary practice, scratching the consultation table is indeed often the best way to draw attention from the cat;
- Visualization of rapid movement triggers approach. Experience nevertheless allows the cat to recognize a motionless prey and to attack it;
- Catch answers to more precise visual and olfactory clues and is triggered by tactile stimuli.

Feral cats hunt every 2 or 3 days, whereas domestic cats can hunt everyday. (©C. Chataignier).

Leyhausen’s studies seemed to indicate that prey capture, killing and consumption were indeed independent actions. Capture and killing appeared to be also independent of hunger but if hunger pre-exists, predatory sequence is complete as hunger is the only reason that explains consumption.

Each stage induces in turn the following one, which permits the succession of all sequences by different stimuli. Hunger is not compulsory to trigger prey seeking but increases kill probability. Satiety does not inhibit sacrifice.

The domestic cat hunts quite often but rarely eats the prey. The system of independent phases proposed by Leyhausen would indeed explain that cats just fed can display all sequences without food intake. Some steps can be repeated and the owner can attend a cruel predatory play, during which prey is still alive while being handled in all ways. The fact of bringing back the prey to the owner can be falsely interpreted as a proof of the maternal like type of bond.

The Most Common Types of Prey

Each catch represents only a small percentage of the cat’s daily energy requirements (the caloric content of a mouse can be estimated at 30 kcal). It is possible to find up to 12 small preys in a feral cat’s stomach. This represents an adaptative behavior to scarcity periods. Continental cats essentially hunt young lagomorphs and rodents. Birds come after in the list and reptiles even lower. Cats living on islands depend mainly on rats, mice and sea birds.

Cats are versatile and generalist hunters. They can easily move from one prey type to another depending upon ecological evolutions. They can even go to some domestic feeding during scarcity periods. On some islands, feline predation has been put as the cause of some species extinction, according to Bateson and Turner (1989). Studies focused on the effect of predation on wildlife are however limited and it is unclear whether the extrapolation of these data to the global feline population is accurate.

Techniques to Limit Predatory Behavior in Domestic Cats

Attempting to decrease predation by house hold cats is probably a worthy goal. Predatory behavior is a normal behavioral pattern in cats but often distressing to their owners. Although 6 of 10 cats (in the United States) and most pure breed cats are kept indoors and therefore cannot hunt, in Europe many cats (7 of 10) have an outdoor access so they can hunt and kill small rodents and birds. Owners often find this behavior objectionable especially when cats direct the behavior toward song birds and/or bring prey home.

Predatory behavior is best prevented by keeping cats indoors and obtaining kittens from queens that do not hunt (so as to get inexperienced individuals). In addition, keeping a quick release cat collar with a large bell on the cat may diminish their proficiency. Nelson et al (2005) compared collar mounted warning devices on reducing predation in cats in the UK and found that there was no significant difference in the prey return rate between cats wearing collars with one bell, two bells or a sonic device.

The cat has been domesticated for nearly 6000 years but has not lost his exceptional hunting skills thanks to the independency of predatory sequences. Cats can easily return to the wild and survive without human intervention.

Domestic Feeding

Cats spend from 1 to 2% of their awake time eating. When feeding a household cat, the food can be provided either in a controlled manner i.e. as meal feeding or as free choice feeding. Regardless of the type of feeding regime chosen, it is useful to establish regular feeding and eating patterns for house hold cats (Table 2).

The Place of the Feeding Bowl

The territorial organization of the cat’s life must be taken into consideration. Each spot has a defined dedication for the cat: feeding, resting, playing, eliminating. And these functions are not mixed. You would not eat in the middle of the train entrance or in your toilets. It is the same for a cat! (Figure 5).

Figure 5. The minimal distance between the feeding place and the elimination and resting places.

Bowls have to be small, to control intake and to encourage frequent refilling. Regular cleaning of the bowl is necessary, to avoid off smells and safety issues. In homes with multiple cats, each cat should be offered their own food bowl. Antagonistic interactions between cats may restrict some individuals from access to food and water bowls leading to weight loss and perhaps medical complications. In addition, because cats may not share space equally, food bowls should be allocated through out the environment, not all in one location. Care should be exercised to note where individual cats spend most of their time and place food and water bowls in those locations. Litter boxes should be placed at a significant distance from the feeding location.

Meal Feeding

When pet owners use a meal feeding method to feed their cats they either control the time the food is provided or the portion size provided. It is the best method for canned food so as to secure freshness and safety. Leaving leftovers for hours in the bowl indeed leads to bacteriological risks and palatability decrease due to organoleptic deterioration. Manufacturers have understood this problem and now propose single portions diets. Because cats eat multiple small meals when hunting, most household cats find single meal feeding unsatisfactory if it is the only method chosen. If an owner chooses to feed its cat in a time controlled manner, at least two meals per day should be provided. However, an increased frequency of meals may help control hunger and decrease excessive food soliciting behavior. It is indeed amazing to see cats learning to detect when they can get extra food. They associate some events to a high probability to get reward e.g., during the advertising break of the evening TV movie or when the owner puts the kitchen in order (they can be warned by the noise of plates in the dishwashing machine!).

Meal Feeding Methods

- With time controlled feeding, the food is left available for a set amount of time and then picked up and not provided again until the next feeding time. For most cats, 30 - 60 minutes should be allotted for eating when fed in a time controlled manner (Case, 2003).
- With portion controlled feeding, the amount of food provided is measured and placed in the bowl and once it is consumed, no more food is provided until the next meal. For a single household cat, portion control can help control weight while potentially allowing the cat to eat several meals through out the day, something that cannot occur with timed feeding.

Meal feeding in either manner has several advantages. It allows owners to determine how much food the cat consumes in a 24 hour period. Food consumption is often a good marker of health and knowing how much the cat eats can help an owner determine how the cat feels. If multiple cats reside in the home it may allow the caregivers to assess each cats eating pattern and access to food and may help the owner to recognize health or social problems in the cats. Meal feeding also allows the pet and the owner to interact several times a day strengthening the human-animal bond.

Free Choice Feeding

In free choice feeding food is provided to the cat at all times so that the cat can eat multiple small meals in a 24 hour period. Domestic cats often eat multiple small meals through out the day. Depending upon observations, this number varies from 8 to 16. This is linked to the evolution of the cat to an opportunistic feeding pattern, nibbling small amounts of food on numerous occasions. This method relies on the cat ability to self regulate intake. This is the best option when the cat is fed with dry food. However, due to the increased palatability of commercial cat foods coupled with reduced exercise, cats may over eat and become obese. In addition, self feeding does not necessarily allow the human caregiver to determine daily intake especially in homes with multiple cats. With self feeding, a change in food consumption and resultant weight loss may not be noticed for some time perhaps imperiling pet health. Self feeding also limits the pet-owner interaction around feeding time.

Some pet owners may combine meal feeding and free choice feeding by providing set meals of small amounts of wet food and provide dry food in free choice. It creates a social re-enforcement of food intake, although this effect is much less important than in dogs. The cat is invited to eat wet food as a "plus" and the calorie content is added to the normal dry food ration spread over the day. In some cases the energy balance can be excessive. The practitioner must keep in mind that some cats may have difficulties to regulate their energy consumption when fed two different types of food: a strict control of the quantities fed to the cat and of the nutritional balance of both types of foods is highly recommended.

The Role of the Human in Feline Feeding

Because most household cats do not hunt to meet their nutritional needs, when they are hungry they target their pre-meal behaviors and food soliciting behaviors towards the humans within the home. These behaviors include vocalization, rubbing on nearby objects and on the humans. Often if the person moves in the direction of where the food is provided or stored, the cat may run in that direction or continue to wind between the legs actually impeding progress. It has been suggested that people who feed the cat have a better relationship with the cat perhaps because of these opportunities to interact (Geering, 1989).

Owners often inadvertently assume all vocalization and attention is in fact a food soliciting behavior. They then respond by feeding the cat, resulting in a potent learning process. The owner’s response is acting as a reward for vocalization and attention seeking behaviors which usually will increase in frequency and intensity. Not only can these behaviors be distressing to the owner, excessive food intake will also lead to obesity and related medical problems. Owners should learn to recognize when enough food has been provided and consumed, and not reward these behaviors with food. They then should find an alternate activity such as play or grooming once the nutritional needs have been met.

Observations of Feeding Behavior: How the Cat Eats

Many nutritional studies are based on the study of the factors affecting the amount of food that cats ingest. The regulation of ingestion is a complex and still poorly understood phenomenon. The frequency and size of meals represent two key parameters of feeding behavior.

Number of Meals Per Day

Each cat has its own way to dispatch its meals throughout the day. A cat generally needs 3 weeks to get a stable life pattern. In an ad libitum situation, it ranges from 3 to 20 meals per day (Kane et al, 1981; Houpt, 2005), like water intake (Mac Donald et al, 1984). In cattery conditions, when cats are fed ad libitum with dry food, the dietary consumption is influenced by the night and day alternance: the dietary consumption at night is often inferior to consumption during the day, but during the night, the meals are larger and longer (Kanes et al, 1981; Royal Canin Research Center: internal data, 2004).

Meal Size, Meal Duration and Speed of Eating

Meal size increases with palatability (especially the first meal) or when the feeder goes from meal feeding to ad libitum feeding. The average duration of a meal is almost 2 minutes (Figure 6). The speed of eating is an important criterion for the owner’s perception. It is in fact much more influenced by food structure than palatability. For dry kibbles, the eating rate increases from 2 to 4 grams per minute. For canned product, it lies between 4 and 8 grams per minute.

Figure 6. The feeding pattern of the cat over 24 hour (Source: Royal Canin Research Center; internal data, 2004).

Studies done at the Royal Canin Research Center illustrate how the number of meals, quantity of ingested food and the speed of ingestion are significantly different for different breeds in exactly the same environmental conditions and sharing the same nutritional history (Table 3). The frequency of the meals is the most variable parameter between the different breeds studied with Bengal cats showing the highest frequency. Maine Coon cats tend to take the largest meals and Persian cats the smallest. Persian cats spend twice as much time per meal compare to the average of the other breeds studied (3’27’’ vs 1’49”) (Royal Canin Research Center: internal data, 2004).

Types of Prehension

Prehension of food explains the apparent eating rate of cats. The cat has a small mouth and small teeth which are better suited to holding and killing prey rather than grinding and chewing food (Case, 2003). The initial role of the canine teeth is to seize prey whereas the carnassials tear flesh.

Innovative studies conducted by Royal Canin in collaboration with the École Nationale Supérieure des Arts et Métiers (ENSAM) in France, demonstrated that cats exhibit three distinct methods of dry food prehension. The most common method is called labial prehension and involves the grasping of kibble using the incisors, without the use of the tongue. The second method is called supralingual prehension, which involves the cat using the dorsal side of the tongue to "lap up" their food. The third method is called sublingual prehension, in which the cat applies the ventral side of the tongue to the kibble, then turning it backwards (Figure 7).

Figure 7. Prehension modes in cats.

By filming cats from below as they eat on glass surfaces, Royal Canin has discovered that certain breeds of cats are more likely to demonstrate one type of prehension style over another. For example, brachycephalic breeds such as the Persian, have difficulty picking up kibble with their teeth, because of the structure of their head and jaw. In 80% of cases, Persians use their tongue to pick up a kibble rather than their lips or teeth. Mastication is somewhat poor in cats. Jaw joint configuration allows only vertical movement. They often break the kibble in one strike, or even immediately swallow in one gulp! This occurs commonly in cats with dental pain, who avoid breaking the kibbles with their teeth, rather they swallow the kibble whole and even vomiting quickly after.

The role of the tongue is very important when speaking about canned food. It acts like a spoon. Its rugose surface easily catches pieces of loaf or chunks.

Each of these parameters must be considered when feeding cats and creating environments conducive to good eating habits.