Animal Behavior for Shelter Veterinarians and Staff. Группа авторов
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The vertical “tail up” is one of the most important visual signals (see Figure 2.1). It largely signals the desire to interact amicably. The cat receiving a “tail up” signal may reciprocate with an approach, followed by touching noses; rubbing of the head, neck, and body along the body of another cat; and tail twining (Cafazzo and Natoli 2009). A similar approach and rubbing behavior sequence may be displayed toward an individual of another species, especially human, or an object near the intended receiver (Turner 2017). A stiff, lashing tail indicates aggression, and a tail tucked under the body signifies fear in a non‐resting cat. Piloerection, or hairs stiffened and standing away from the body, indicates strong arousal and is usually accompanied by an upright or arched body, but the tail may be erect or low. A tail wrapped around the body of a sitting cat is thought to communicate ambivalence—the cat is unlikely to show aggression but is not enthusiastic about the interaction (Leyhausen 1979).
Figure 2.1 Tail up greeting display.
Source: Reproduced with permission from N. Drain. © Natasha Drain.
Facial expressions are associated with a range of emotions. A distressed cat’s eyes may blink and pupils dilate, ears lower or flatten to the head, and whiskers move forward. These expressions may be accompanied by hissing or other defensive vocalizations (see Figure 2.2). A very relaxed cat is often lying lateral with the eyes, pupils, and ears in a neutral position. A cat resting in a sternal or sphinx‐like position, often with the tail wrapped around the body, may be slightly more anxious or vigilant about the surroundings (Leyhausen 1979; Bennett et al. 2017; Gourkow et al. 2014).
Figure 2.2 Defensive position.
Source: Reproduced with permission from K. Watts. © Katie Watts.
General Appendix B describes common feline body language characteristics.
2.5 Behavioral Development
As a species, cats are extremely adaptable to various environments, but the degree of plasticity is determined by a combination of genetics, prenatal environment, and postnatal experiences, especially those occurring during the socialization development period. Born blind, deaf, and completely dependent on the queen for nourishment, the neurological, musculoskeletal, and cognitive maturation of a kitten from birth to adulthood is a short but complex process.
2.5.1 Genetics
Comparative genetics studies have revealed the domestic cat varies from wildcats at 13 chromosomal loci, many of which code for genes related to neurodevelopment or neurotransmitters known to affect various emotional or motivational states, perhaps revealing the genetic basis for tameness (Montague et al. 2014). Humans have a long history of enhancing certain traits through genetic selection, and this is most evident in the domestic dog, the species with the greatest morphologic diversity on earth. Humans have a much shorter history of selectively breeding domestic cats, and the primary objective has been advancement of desired physical, not functional, traits. Nevertheless, consistent breed predispositions for certain behavioral characteristics seem to exist. Several observational and survey studies have identified differences among purebred cats in terms of aggression, propensity to elimination outside the litterbox, playfulness, shyness, and activity level (Mendl and Harcourt 2000; Wilhelmy et al. 2016; Salonen et al. 2019). Many people believe that coat color and certain personality traits are linked. For example, calico or tortoiseshell color cats are often thought to be more aggressive and anxious (Stelow et al. 2015). While the owner‐queried survey results of Stelow et al. (2015) did indicate some minor increases in aggressive behavior within sex‐linked orange color cats, Wilhelmy et al. (2016) found that differences in behavior among purebred cat coat colors were largely explained by breed alone.
Cat personalities seem to be stable and vary along several dimensions of confidence, nervousness, sociability, and activity (Lowe and Bradshaw 2001; Karsten et al. 2017; Litchfield et al. 2017). Personality, like most behavioral phenotypes, is a confluence of environmental and genetic influences. Kittens sired by males with outgoing and friendly personalities were found to show more approach and affiliative behaviors to people, be less stressed by the approach of unfamiliar people, and be more likely to spend time near a novel object (McCune 1995; Reisner et al. 1994). However, handling and socialization provided a protective effect against some fearful and defensive behaviors. Friendly sired but unsocialized kittens behaved in a similar manner to unfriendly sired socialized kittens (McCune 1995).
2.5.2 Sensitive Periods of Development
2.5.2.1 Prenatal
The nutritional status of a dam and her exposure to certain stimuli during the 63‐day gestation period can influence postnatal behavior of her offspring. For instance, kittens show a preference for certain flavors fed to their queen during the prenatal period (Becques et al. 2009). Queens placed on protein‐ or calorie‐restricted diets can produce offspring with elevated emotional reactivity and impairments in social interactions, environmental exploration, and learning (Gallo et al. 1980). Even food restriction limited to the second half of gestation can result in abnormal physical and emotional traits (Smith and Jansen 1977). Studies from other mammalian species indicate offspring born to dams exposed to excessive environmental stressors, such as unpredictable noise stimuli, may suffer from impaired cognitive and neurologic development (Schneider and Moore 2000). Kittens exposed to significant prenatal stress may suffer similar developmental dysfunction.
2.5.2.2 Neonatal (0–14 days)
The neonatal period consists of the first two postnatal weeks. The queen encircles the kittens with her body and legs immediately after the birth of all of the kittens. The kittens are suckling within an hour of birth, and a loose teat order is established by 12 hours post‐parturition (Ewer 1960; Houpt 2018). The kittens engage in little other than suckling and sleeping during the neonatal stage. Waste elimination is initiated by the queen via grooming of the perineal region. Neonate kittens do not have the ability to thermoregulate, and thus body temperature is maintained by huddling with their littermates and mother (Jensen et al. 1980; Olmstead et al. 1979). Tactile, thermal, and olfactory stimuli help the kitten orient to the queen and littermates, as the eyes and ears are closed during much of the neonatal period. Orienting to the queen’s abdomen and suckling are highly reflexive behaviors (Raihani et al. 2009), although nursing is largely initiated by the queen during the first two weeks. Purring is observed in nursing kittens and may serve to communicate active suckling to the mother (Bradshaw 2017). The first set of teeth erupt between 2 and 5 weeks of age with adult teeth erupting at approximately 12 weeks. Movement at this age occurs by limb paddling or pulling of the body by the front limbs due to weaker neuromusculature of the hindlimbs in this early stage.
Kittens are usually able to hear by the fifth day of life, although the external pinnae do not become erect and the ear canal does