Early Phase

During the first 3 weeks of life, feeding sessions are initiated primarily by the queen. She arouses the usually sleeping kittens by her movements and licking and positions herself around the young, almost enclosing them with her limbs and ventrum. The rooting reflex causes the kitten to burrow into a warm object, usually its queen or littermates, and may be present up to 11 days (see Figure 2-9). Maturation of the senses, particularly vision, and of homeostatic mechanisms alters all neonatal reflexes. During the appetitive searching component of nursing behavior, the kitten’s movements toward the queen are awkward, but they become progressively more coordinated as the kitten’s muscular and nervous system control increases with age. Communal nests allow some queens to actually have nursing coalitions.¹²⁴

The sucking reflex is present at birth and can be tactually stimulated over a large perioral area.¹¹⁷ Such stimulation results in a head turn in the direction of the stimulus and accompanying sucking movements (see Figure 2-10). Small objects in the mouth also stimulate the reflex, which is strongest immediately after awakening. Within a few days, the sucking reflex is limited to lip contact, and foreign objects in the mouth are rejected.¹¹⁷ That is probably when olfactory cues become more important in teat location than trial and error, the initial method the kitten uses. Destruction of the olfactory bulbs produces kittens that cannot initiate sucking even with previous experience, but it does not interfere with their ability to learn how to suck from a bottle.¹¹⁰ This is also consistent with the finding that odor cues may be important to locate the mother, and tactile cues help locate the nipple.¹¹⁸ The sucking reflex normally disappears by 23 days.^108,110 Taste and texture can be discriminated shortly after birth but are apparently of minor significance in normal sucking.¹⁵³

The kitten’s initial contact with the teat results in a withdrawal motion of the head, followed by a downward motion and perhaps another withdrawal before the teat is finally enclosed by the mouth. This random head bob decreases as the kitten’s ability to discriminate the teat increases. By 4 days, kittens are relatively proficient at suckling the queen, and by 1 week, little nuzzling is necessary.

Before the end of the third day and sometimes as early as the first, a form of teat preference develops, which may become very rigid. Rear nipples are used preferentially.⁵² In large litters there is an initial competitive scrambling and pushing, which decrease as the preference for specific teats develops. As many as 80% of the individuals nurse only from a specific nipple.^61,160,166 When preferences are more loosely defined, a kitten nurses from a pair of teats, usually near each other, or from a general region of the mammary surface.^160,161 Occasionally individuals do not develop any preference and will nurse randomly instead.^159,166,167 The incidence of teat preference is independent of litter size.¹⁶⁰ Development of teat preference may serve to minimize claw injuries among littermates, provide optimal stimulation to ensure milk production, and allow more rapid completion of nursing. Variability has developed with domestication because the behavior has not been considered in selective breeding.

If a specific teat is not sucked for about 3 days, milk production by that mammary gland stops.⁶⁵ The appearance of the mammary region of a nursing queen makes it obvious that only certain nipples are used.

Nursing positions are identified by olfactory means, and kittens seldom nurse strange teats after the first few days if normal nursing is allowed.^62,71 A kitten that takes the wrong nipple may leave it immediately or may continue to suckle until the rightful “owner” nuzzles at it. Only when nursing the preferred teat will a kitten tenaciously cling to it when challenged.¹⁶⁰ Unlike pigs, kittens nursing from the cranially positioned teats do not necessarily become larger than their other littermates.⁶¹ Teat positions are maintained for approximately 1 month, at which time the preference becomes less rigid. At about the same time, kittens show less discrimination in nursing foster queens.^62,64 In cat colonies where multiple litters are housed together, kittens commonly suckle queens indiscriminately, with little or no teat preference development. When bottle feeding is necessary, repeated use of the same bottle and nipple and avoidance of noxious or unfamiliar odors on human hands are desirable.

The “milk tread,” usually considered a part of the nursing behavior of kittens, consists of rhythmic alternate movements of the forepaws against the mammary gland. Initially, a somewhat similar pattern is observed as the neonate tries to steady its suckling position. Functionally the milk tread may serve to push the queen’s skin away from the neonate’s nose but soon becomes more important to help stimulate milk flow.^52,63,64 The behavior is primarily used when milk is not coming as fast as the kitten can drink and is usually alternated with nonrhythmic sucking.

Purring by both the queen and the kittens often accompanies nursing.²⁸

The neonate initially spends a great deal of time nursing. Each session may last up to 45 minutes, totaling about 8 hours per day.^13,52,162 This amount of nursing allows most kittens to double their birth weight in 1 week, triple it in the second week, and quadruple it by the end of the third week.^13,183 Milk production does increase for large litters but not in direct proportion to litter size.⁵² In addition, there is as much as a tenfold increase in subcutaneous body fat for insulation to aid in homeostasis.¹⁸³ The mean growth rate for the first 8 weeks is 7.3 g/day (kittens from litters of seven to eight) to 13.7 g/day (kittens from litters of two).⁵²

Intermediate Phase

During the second phase of nursing behavior, suckling is initiated about equally by the queen and the kittens. At first, 40% of the feedings are kitten initiated, with marked increases later.¹⁵⁹ By 4 weeks, approximately half of the contacts are initiated by the kittens and half by the queen.⁵² The intermediate phase generally lasts from the second or third week of life to the end of the fourth or fifth week, when development of vision, use of visual cues, and coordination of motor skills allow the kittens to actively approach the queen even when she is not in the nest area. She in turn lies in lateral recumbency to present the mammary region for the young to nurse (see Figure 6-9). Toward the end of this phase the kittens may initiate sucking with the female in a standing position, but she will then respond by lying down.

Avoidance Phase

Near the end of the first month, kittens are very active in initiating nursing and the queen becomes increasingly evasive of them—the avoidance phase. By 8 weeks, only 5% of the contacts are initiated by the queen, 42% by the kittens, and 42% mutually.⁵² The increasing demand for milk at this time already has the mother eating twice what she normally would to keep up.¹²⁸ At first the female avoids some of the advances of the young by jumping onto objects they cannot reach, but as the kittens’ motor skills develop, they become increasingly difficult to avoid. Avoidance is made even more difficult because their efforts are usually individualized rather than coordinated. Her tactics will change to making nursing progressively more difficult. The queen may lie in sternal recumbency to prevent access to the teats, and if the kittens become too persistent, she bats at them or gets up and moves away. When she does allow nursing, the queen is usually sitting or standing, with the kittens contorted into several postures. If she has only one kitten in the litter, she may not show this avoidance phase.

Weaning Phase

Weaning is a gradual, variable process that results from inaccessibility of the queen and increasing ability of the kittens to hunt. In the wild, cats do not wean their young, but if the queen is still lactating, the milk is of little nutritional value after 12 weeks.^35,169 When this decreased value is coupled with normal dispersion necessary to obtain prey, weaning simply happens. Even without increased avoidance by the female, there is a natural tendency for the young to decrease their nursing at about the time of normal weaning.⁷⁹ When kittens are confined with the queen or have ready access to her, nursing may continue for several months, although 8 to 10 weeks is most common.^71,109,160 There is an inverse correlation between litter size and length of time the female allows nursing.¹³ Smaller litters nurse longer. Kittens born in the spring often continue to nurse for 3 to 5 months, until fall.^95,185 Some queens will let a kitten nurse until just before the birth of her next litter, as much as a year later, then aggressively chase it away. The older offspring may return shortly after she queens and continue nursing, lying among the neonates.

Developmental Hunting

Rudimentary predatory behaviors can be seen at 3 weeks of age.¹⁴³ Then, prey recognition begins toward the end of the first month, when the queen brings home dead prey for her offspring to smell and eventually taste. In another 3 or 4 weeks, the queen brings weak or wounded prey so the young can begin acquiring their own hunting skills. For each of these behaviors, the queen’s own prey-killing and eating instinct must be inhibited, primarily in response to kitten-related cues. Kittens will interact with prey more quickly and for longer periods immediately after the mother’s bout.³⁹ Between 4.5 and 7 weeks, most predatory behaviors are significantly correlated with each other.

Hunting techniques are developed by practice during play and during prey sessions with the queen. The chase and catch aspects of hunting are thought to be innate, but manipulative and positive contact responses are acquired behaviors.^13,72,158 The chasing, catching, and killing aspects of hunting may also be learned or at least perfected by learning. The nape bite is the prey-killing behavior that does not usually appear in play and must be perfected with practice on actual prey. When the kitten’s hunting style fails and the mouse or other prey escapes, the kitten is given another opportunity because the queen recaptures the mouse and drags it back to the young. Kittens follow the queen on hunting trips to perfect their skills as they approach self-sufficiency. By 15 to 18 weeks, exploration and hunting take up to 68% of the young cat’s time.¹⁸²

Predatory tendencies are inherited and have probably been modified by domestication, making prey-killing cats generally less defensive and more aggressive than nonhunters.¹ These tendencies are influenced by early experience. Competency in predation is related to the presence of the queen during the kitten’s exposure to prey and to experience as an adult.^23,37,38,87 Kittens hunt the same prey they observed their mother kill and bring home. Cats are better able to catch prey if they had experience with prey by approaching, manipulating, and biting them in a play context as kittens.³⁶ Learning to kill prey is generally limited from the sixth to approximately the twentieth week, but for the few cats that learn after that age, the process appears almost laborious.¹²³ If the young never observe this process, approximately half will learn hunting skills on their own. Even if a kitten is never exposed to hunting, at an appropriate age the cat responds to the squeak of a mouse by attacking it.⁷⁴ The way a kitten is raised will also be important in future hunting behavior. A young kitten raised only with a mouse or rat seldom becomes a hunter, but the approximately 17% that become hunters do not generally hunt the type of prey with which they were raised. If the kitten is raised with a mouse or rat in addition to other kittens, socialization to the natural prey is minimal.^111,112

The Hunt

Prey capture

Although rustlings or squeaking may initially attract the cat’s attention, the sight of moving prey is the primary factor initiating hunting behavior. With experience a cat can learn to recognize immobile prey.¹⁷⁶ Kittens can visually orient to prey as early as 9 days of age.¹⁵⁸ Hunger and prey killing are independent, but the former can lower the threshold for the latter.^20,122,138 After being alerted to potential prey, even prey as large as itself, a cat approaches the prey in a stalking ambush. Initially the cat will head to a particular area, and when it arrives, the gait changes to a slow walk as it looks over the area.¹⁷⁶ Once prey has been spotted, the cat crouches close to the ground and uses a slinking trot, stopping periodically behind cover. It lies there temporarily with forelimbs under and elbows protruding. The entire pes is on the ground under the cat. The head is stretched forward, as are the ears. Whiskers are spread. As its eyes follow the prey, the cat usually twitches its tail.¹³ The stalking continues periodically until the cat nears striking range. At that time it again stops and lowers its body, proceeding at a cautious, low-profile walk (Figure 7-1). At the last available cover, there is a final pause during which its tail twitches with great intensity and the caudally positioned pelvic limbs tread, swinging its entire hindquarters to and fro.¹³ Then, after a short run, if necessary, the cat springs forward to seize the prey. In this pouncing attack, its pelvic limbs usually remain on the ground and only its forefeet are used in the capture (Figure 7-2). As soon as its forelimbs leave the ground, its rear limbs spread apart to stabilize the cat and help it brake. In this way, the cat can change direction to accommodate zigzags made by the fleeing prey. If instead both thoracic and pelvic limbs were off the ground during the lunge, the cat would be committed in its direction of travel. The cat that misses its intended prey makes no attempt to correct the error. Instead it temporarily withdraws, and if conditions are still favorable, it renews its attack.¹²³ On average, a cat will make 3.6 pounces for each vertebrate it catches while hunting.¹⁴⁶ Cats have been known to leave one mouse in favor of another more active one.¹³

Figure 7-1 The stalking walk of prey hunting.

Figure 7-2 The pounce used in prey capture.

Nonrodent Prey Capture

Studies of ingested material in feral cat populations confirm that mice are the cat’s primary food. Other prey varies seasonally but includes voles, bats, small reptiles, carrion, rabbits, birds, and insects. Rats weighing less than 200 g are another prey species, but half are killed and not eaten.⁴⁵ Feral cats tend to be opportunists and scavengers, so the type of prey is largely dependent on its relative abundance.⁴⁹ The availability of commercial food or table scraps does not decrease consumption of wild prey.^51,81

Birds do not make up as much of the cat’s diet as is commonly believed, with human activity being 56 times more likely to kill birds than cats.^148,149 Spring and summer hatches provide most of the supply, and cats that hunt near roads apparently kill more birds than those hunting in fields.^58,132 Most garden-type birds tend to breed at the edges of woods and in clearings where they would be preyed upon by several species of animals, not just cats.¹³³ Special hunting skills are required to catch flying prey, so not all cats are successful. The best hunters attack as soon as they spot the bird because ambush techniques often send the intended prey hopping or flying. The bird is usually caught with one or both forepaws and instantly brought to the mouth for the nape bite. If the bird is long necked, the nape bite may be directed close to the head or the shoulder. Instinctively, the latter is more commonly used because the size disparity is greater between body and neck than between head and neck. The best hunters do not release the catch for a better neck bite, because the bird tends to escape. Except for a few feathers, which may be plucked, the entire animal is usually eaten, beginning with the head.^13,28

Once a cat starts hunting birds, it is almost impossible to prevent its continuation. At best, throwing things at the cat causes it to hunt on the sly. To avoid a bird-catching cat, a kitten should be selected whose mother does not hunt birds, or it should be adopted at about 6 weeks of age. In both cases, the kitten should be confined and not allowed exposure to birds, at least until after it is a year old.^27,84 Placing a bell on the cat may give birds enough of a warning.²⁷

Rabbits caught are usually juveniles, probably because full-grown ones are too powerful.¹⁷⁶ Even then, the difficulty of the catch of such a large animal is still greater than that of other species.¹⁷⁶

Another less common prey for the cat is the fish. Although most cats leave fish alone, a few are noted for pursuit of the family’s goldfish. The fishing cat of Malaysia is very adept at hooking fish with its forepaws, as are certain individuals living in coastal regions or near lakes and streams. Fishing is one of the earliest skills for which selection was made in domestication. Like good bird hunters, good fishers do not let the catch go to obtain a better neck bite, because that leads to the fish’s escape. Cats having no early fishing experience may actually run away from a fish flopping along a shore.

In certain parts of the world, livestock injuries from vampire bats can be significantly reduced by hunting cats.⁵⁴ Successful catches tend to occur when the bat is distracted during its hunting approach, while defending itself from the cattle, or when weighed down after feeding.