section epub:type=”chapter” id=”c0013″ role=”doc-chapter”> Jacqueline Mary Ley and Kersti Seksel A cat’s behavior is the result of a complex interplay of genetic predisposition, what the cat has learned from previous experiences (good, bad, and neutral), and the cat’s current environment. Although some behavior patterns are common to all members of a species, others are unique to the individual. It is essential to understand normal cat behavior to assess behaviors that owners are concerned about. This will help the veterinarian determine whether the behavior is normal and adaptive or abnormal and maladaptive. Cat; feline; behavior; adaptive; maladaptive; vision; hearing; olfaction; vomeronasal organ; flehmen; body language; felinine; learning; allogrooming; social organization; time budget To understand the behavior of cats, the veterinarian must first look at the physical characteristics of the cat, such as its size and sensory capabilities, because these are intertwined with behavior. Only by appreciating the behavioral biology of the domestic cat is it possible to understand their behavioral needs. The domestic cat is a small, crepuscular, solitary hunter of the felid family. Whether the domestic cat is a unique species or a subtype of the wild cat (Felis silvestris) of northern Africa remains controversial.1 The cat evolved in arid areas and hunts small animals such as rodents, frogs, birds, and reptiles. They are small, tending to weigh between 2 kg (4.4 lb) and 8 kg (17.6 lb) and have large, forward-facing eyes; large, mobile ears; and sensitive vibrissae on the face that aid in detecting prey in dim light. They have large, ventrally flattened canine teeth and sharp retractable claws on all toes to catch, hold, and kill prey. The cat is an ambush hunter. It locates prey using its sensitive hearing, vision, and sense of smell. It then stalks the prey silently until it is close enough for a sudden rush and grab. Cats do not possess the stamina to chase prey for prolonged periods of time. However, they can climb and jump up to five times their own height.1 Being small, they are potential prey for larger animals, so their agility is an advantage not only for hunting but also for escaping when they are being hunted. One of the reasons that cats are so appealing to people is their large, prominent eyes. Large eyes are necessary for seeing (and hunting) in dim light. Cats’ eyes have many characteristics to maximize the visual field and collect light entering the eye to stimulate the retinal cells.2 The cornea is large and bulges outward, which allows about five times more light to enter the eye than does the human cornea.1 The retina has approximately 25 light-sensitive rod cells for every color-sensitive cone cell. When rod cells in a cluster are stimulated by light, they all stimulate one nerve fiber. This results in cats being able to see in very dim light, albeit a fuzzy image.1 The tapetum lucidum under the retina reflects light back to maximize the chance of rods being stimulated. This layer is what makes cat eyes glow yellowish green when light shines into them. Cats have little need for color vision because they hunt mainly at night, and, like most prey species, non-purebred cats do not have a wide range of coat colors. It appears that cats can see yellow and blue wavelengths of light and can be taught to distinguish among red and other colors. However, this is difficult for them to learn, which suggests that cats are just not interested in colors.1 The lens of the eye has a limited capacity for accommodation. This means that a cat’s vision is best at approximately 2 to 6 meters (6.5 to 19.7 feet) from the viewed object.1 This is why cats may have trouble taking treats from an owner’s hand. To maximize visual acuity, they have multifocal lenses that focus light at particular wavelengths. The slit pupil prevents the loss of visual fields that can focus at set wavelengths and maximizes the cat’s vision.3 Binocular vision aids the cat in judging distances for catching prey, climbing, and jumping. The binocular overlap is about 98 degrees, which allows cats to judge distances very accurately.1,2 Their accuracy is even more amazing considering how short-sighted they are. Cats are very attuned to even small movements in their visual field. The eyes of cats are not functional at birth. The eyelids open between days 14 and 21. Vision develops with experience. If kittens are deprived of vision through blindfolding before their eyes open4 or are housed in environments that are altered to show no horizontal lines, the kittens do not develop normal vision, even though the eyes are structurally and functionally normal.5,6 A cat’s large, mobile pinnae collect and funnel sounds into the ear canal. Each ear can move independently of the other, and the ears can swivel almost 180 degrees, effectively giving them ‘surround sound.’ When tracking a sound, such as that of a prey animal, cats use a combination of the interaural time differences for sounds to reach both pinnae, level differences between the pinnae, and directional amplification effects of the pinnae to localize the sound and orientate their head.7 Cats have a well-developed sense of smell at birth. These nerves are myelinated at birth, in contrast to most other neurons in the nervous system. This allows signals to pass rapidly to the brain. The kittens use their sense of smell and touch to find the queen’s teats. If they are unable to smell (e.g., because of an upper respiratory tract infection), kittens cannot find the queen’s teats and feed.8 Cats use their sense of smell for locating prey and evaluating communication signals left by other cats. Odors play an important role in the social organization of cats and in reproduction. The feline nasal mucosa is between 20 and 40 square centimeters in area, small when compared with dogs, although it still eclipses the human nasal epithelium. To further aid in scent detection, the cat has two structures: the subethmoid shelf1 and the vomeronasal organ (VMO).9 The subethmoid shelf traps air and scent particles taken into the nasal cavity allowing more time for them to stimulate receptors in the olfactory mucosa.10,11 The VMO sits between the oral cavity and the nasal cavity. It has connections with the nasal cavity and the oral cavity. The receptors of the VMO are different from those of the nasal epithelium. The gape or flehmen response may be performed after the cat has sniffed or even licked at a scent source. By wrinkling the upper lip and opening the mouth, the cat opens the ducts of the VMO and pumps saliva and the scent into the VMO.9 Cats cannot fully evert their upper lip as horses and cattle can because of the frenulum between the upper lip and upper jaw. The gape or flehmen response is seen when tomcats encounter urine from another cat. However, queens and neutered cats also exhibit this behavior when investigating odors. Anyone who has pet a cat knows how important physical contact is to cats. Touch is used to build social bonds within feline social groups. The response of cats to touch and temperature varies across their bodies. Cats do not react to heat on their bodies until the temperatures reach 51° to 54°C (124° to 129 °F). However, the skin around the nasal area is exquisitely sensitive to temperature changes, reacting to temperature increases of 0.2°C and decreases of 0.5°C. This ability is an advantage for locating prey. Cats have individual differences in their preferences regarding petting and handling. Some like very strong pressure, whereas others prefer a light touch. Cats have specialized tactile vibrissae on their faces and forelegs. The vibrissae are long, thick hairs that are obvious against the coat of the cat. They sit in a large follicle with a sebaceous gland attached. Striated muscle attached to the follicle allows the vibrissae to be voluntarily moved. The follicle has several nerve receptors associated with it. These are sensitive to pressures on the vibrissae as light as 2 mg or displacement of the whisker by as much as 5 Angstrom (or 0.0005 micrometers),12 and they are sensitive to movement of the vibrissae from the normal position. The facial vibrissae, better known as the whiskers, are synonymous with cats. These are arranged in rows on the upper lips. The upper rows move independently of the lower rows. Cats fold the whiskers back when relaxed and spread them when walking or showing interest in something. Because cats cannot see things that are close, their whiskers are important for the location of prey, food, water, and other objects close to the face. Whiskers also aid in spatial awareness. They are also used to locate prey at the final pounce, especially in dim light. Cats have a superciliary tuft above each eye and two tufts between the ear and point of mandible known as genal tufts 1 and 2. Genal tuft 1 is dorsal to genal tuft 2. These vibrissae also help in spatial awareness. There are also vibrissae on the back of both carpi just dorsal to the accessory pad. It is thought that these tufts aid the cat in using its forelimbs for activities such as hunting. The sense of taste is important to cats. They have two types of taste buds on their tongues: mushroom-shaped papillae at the front and sides of the tongue and cup-shaped papillae at the back of the tongue. Cats can taste salty, bitter, and acid. They have little reaction to sucrose and tend to drink sweet water only if the sugar is masked by salt. In fact, cats lack the ability to taste sweetness, unlike other mammals. The taste receptor for sweetness is made up of two proteins generated by the genes Tas1r2 and Tas1r3. In cats the Tas1r2 gene does not code for the normal mammalian protein, thereby impairing the function of taste receptors for sweetness.13 Cats send signals using body language—that is, by changing their posture, the position of their limbs and ears, and the size of their pupils and by puffing up their fur to appear larger. Cats are very expressive, and it can help when learning cat communication signals to look at each area of the body separately. One method cats use to send messages to other cats, other animals, and humans is by using their bodies. The size and shape of the body, position of the ears, size of the pupils, size and position of the tail, and visibility of weapons such as teeth all convey important messages to others. In general terms a confident cat stands tall and evenly on all four feet, with its tail up or level with its back and its ears facing forward. An attacking cat usually makes itself appear larger by standing at its full height and bristling its hair coat to appear larger. The tail will also be raised, with its fur puffed out. When a cat really wants to convey a message to an opponent that it is ready to fight if the other does not back down, the cat will arch its back (Fig. 13.1). The more fearful a cat is feeling, the lower its body gets to the ground. An uncertain cat may take the middle road, often lowering its rump while keeping its forelegs available for striking.
Normal Behavior of Cats
Abstract
Keywords
INTRODUCTION
THE BIOLOGY OF CATS
SENSE ORGANS
Vision
Hearing
Olfaction
Touch
Taste
COMMUNICATION
Body Language
Body
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