Types of Animal Behavior

The behavior of animals has been broadly classified and studied extensively under categories such as feeding, sexual, maternal, and social behaviors. These studies have helped us to better understand the animals and address their behavioral problems.

27.5.1 Feeding Behavior

Feeding behavior is a circadian rhythm and an important sign of health which is influenced by a variety of internal and external factors. The factors can act on a short-term or long­term basis or be operational only in emergency situations.

Short-Term Control

Taste can either stimulate or inhibit intake. Gastric factors, particularly gastric fill, can suppress feeding whereas the hormone ghrelin can stimulate it. The changes within the gastrointestinal tract such as an increase in osmotic pressure and release of the hormone cholecystokinin cause satiety thus inhibiting intake.

Long-Term Control

The long-term control of feeding in which intake is controlled as part of the regulation of body weight or body fats is especially important in pigs and other domestic animals. The feedback from fat cells to the brain is by leptin which acts on the brain, and as a result, feeding signals are inhibited and feed intake decreases. All these signals are integrated into the brain, involving a variety of neurotransmitters and anatomical sites. The central nervous system controls of feeding are complex. Neuropeptide Y (NPY) and agouti- related protein (ARP) stimulate feeding, whereas cocaine and amphetamine-related transcript (CART) and melanocyte stimulating hormone (MSH) suppress feeding by their actions in the hypothalamus. In addition, when the sympa­thetic nervous system is stimulated, lipolysis occurs and decreases fat stores. Another feedback system is peptide YY (PYY), a peptide released from intestinal cells when ingested fat enters the cells, and suppresses feeding.

Emergency Control

Lack of metabolizable glucose induces eating in response as an emergency mechanism; however, increasing blood levels of glucose does not suppress feeding.

27.5.1.1 Feeding Behavior in Ruminants

27.5.1.1.1 Ingestive Behavior in Cattle

Cattle eat most of their meals during daylight but may graze at night in hot weather. Grazing time is 400-650 min/day at a bite (prehension) rate of 60 per min. Cattle will also eat more in groups than they will individually, and first-calf heifers will eat more if grouped with older cows. Another effect of the social hierarchy during group feeding of grains is that subordinate cows eat faster than dominant ones, probably because they have less time to eat as they are more likely to be displaced by a dominant cow.

27.5.1.1.2 Grazing Behavior

The preferences of ruminants for plant species vary and depend on various factors including the growth stage of the plant (ruminants prefer fast-growing, succulent species). Cat­tle wrap their tongues around grass and pull to prehend it. This method of foraging limits them to plants that are higher than 10 mm. Cattle eat less when environmental temperatures are high. In addition, higher temperature also effects plant growth that may render them less palatable. Food intake falls in estrous cattle.

27.5.1.1.3 Ruminating Behavior

The cows spend nearly 7-10 h ruminating. The composition of the diet especially neutral digestive fiber and the particle size influence the rumination behavior. Apart from the die­tary factors, rumination behavior can be altered by the state of the cow and management strategy. For example, the onset of estrus reduces daily rumination times and the diurnal pattern of rumination.

One of the most common abnormal behavior is the fat cow syndrome wherein the dry cows may overeat, gain weight, and later at calving or during lactation become ill or even die.

27.5.1.2 FeedingBehaviorinSheep

Sheep grazes up to 12 h/day, and most of this time is spent biting.

27.5.1.3 FeedingBehaviorinGoats

Goats eat 12 meals a day with most of it during the daytime. Goats are more selective feeders than sheep and have a longer vertical reach than sheep of the same weight. Browsing behavior is a skill that they learn. Goats learn to break twigs off the plants rather than to chew them off.

27.5.1.4 Feeding Behavior in Dogs

Dogs that have free access to food throughout the day eat many small meals a day, mainly during daylight hours and once-a-day feeding is not “natural,” or not much preferred, by dogs. Addition of another animal to the household increases the original pet’s interest in food. During estrus, bitches tend to eat less. Both cats and dogs have a lower metabolic rate following castration or spaying, which also accounts for their tendency to gain weight.

The common behavioural problems encountered are obe­sity, pica, grass eating, coprophagia, and psychogenic polyphagia.

27.5.1.5 Feeding Behavior in Cats

Similar to dogs, cats eat many small meals (12) per day when given free access to food, but unlike dogs, cats eat both in the light and in the dark. A feral cat with good hunting skills might easily catch 12 mice (or 3 rats) per day. Cats have neither been shown to increase their food intake when housed in groups rather than individually, nor does the sight of one cat eating stimulate other cats to eat when food is offered.

The common behavioral problems reported are wood chewing, pica, obesity, anorexia, and plant eating.

27.5.1.6 Feeding Behavior in Pigs

Pigs are essentially diurnal animals; therefore, most feeding takes place during the day.

27.5.1.7 Feeding Behavior in Horses

In free range, horses normally graze continuously for several hours and then rest, depending on the weather conditions and distances that have to be traveled to obtain water and suffi­cient forage. Grazing behavior is selective to the species it prefers to ingest. Horses show dynamic averaging in which they return to the last patch they grazed after a short absence. Horses prehend at a rate of 25 bites/min. Intake rate depends on bite size and rate. Horses sometimes exhibit pica, includ­ing geophagia (especially the soil rich in iron and copper) and ingestion of stones.

27.5.2 Sexual Behavior

Sexual behavior is important in all species of animals. Sexual behavior includes proceptive and receptive behavior by the female and courting and mate guarding by the male, as well as actual copulation.

27.5.2.1 Physiological Bases of Sexual Behavior

Adult sexual behavior depends on a variety of factors for its expression: These factors are the sex of the animal, perinatal organization, action of hormones, past social and sexual experience, the attractiveness of the potential mate, and the external environment. Both in males and females, the hypothalamic-pituitary axis is involved in the exhibition of sexual behavior. Testosterone acts upon the anterior hypothalamus-preoptic area in conjunction with appropriate stimuli from an estrous female to produce male sexual behavior.

27.5.2.2 Role of Pheromones in Sexual Behavior Manifestation

Pheromones are volatile chemicals secreted/excreted outside the animal’s body to trigger a social response in the members of the same species.

27.5.2.3 Cattle

27.5.2.3.1 Cow

Cow is a nonseasonal, continuously cycling breeder. The onset of estrus occurs more often in the evening and ceases in the morning. Actual sexual receptivity lasts 13-14 h. The estrous cow exhibits increased motor activity and a decrease in feed intake. The more active a cow is, the higher her fertility. Investigative behavior such as flehmen, sniffing, rubbing, and licking increase as does premounting behavior such as standing behind the cow and resting the chin on the back of another, usually another estrous cow. The cow bel­lows frequently, switches her tail and raises or deviates it to one side, and urinates frequently. If a bull is available, the cow tends to approach it. Estrogen levels peak when the cow stands to be mounted. The cow that mounts is usually pre­ovulatory. The mounting cows are also usually dominant over the mounted cows. This represents an interaction between hormonally mediated behavior and social influences.

Visual cues such as homosexual mounting are received over greater distances than olfactory cues and may attract bulls which are separated from the cows.

Silent heats and Nymphomania are commonly reported sexual behavior problems.

27.5.2.3.2 Courtship Behavior in Cattle

The courtship sequence is actually a series of reciprocal interactions between male and female. Starting late in the cow’s proestrus, the bull begins to graze beside the cow, guarding her from any other cattle. His attempts to mount will be repulsed by the cow. During proestrus, most females are attractive to the male and attracted to him but not yet receptive. The bull may attempt to herd the cow away from the rest of the herd. Periodically, the bull will smell and lick the cow’s vulva, often followed by the flehmen response. Some, but not all, dairy bulls flehmen to estrous urine. Fleh- men is followed by an increase in circulating LH. When the cow is in full estrus, the bull will have a partial erection while guarding her, and accessory gland fluid or precoital discharge will drip from the penis. The bull frequently nudges the female’s flanks and may rest his head across the cow’s back while they stand in a T-position. He makes several mounting attempts with a partial erection before the female will stand for him. When she is ready, the cow remains immobile and the bull mounts immediately. He fixes his forelegs just cranial to the pelvis of the female as he straddles her. Ejaculation occurs within seconds of intromission and is noted by a marked, generalized muscular contraction. The bull’s rear legs may be brought off the ground during this act. Dismounting and retraction of the penis follow immediately. The stimulation of sexual behavior caused by a new female is called the Coolidge effect. The Coolidge effect occurs in bulls so that the mounts with intromission and decreased mounting intervals occur when a novel female is provided hourly.

Commonly noted problems among bulls include mastur­bation and impotence.

27.5.2.4 Sheep

27.5.2.4.1 Ewe

Sheep are short-day breeders, showing breeding activity in the winter months as the photoperiod decreases. The ewe is polyoestrous and will cycle several times during one breed­ing season if not bred; the average cycle for the ewe is 16 days (range 14-20). The actual period of estrous receptiv­ity is 30-36 h in the ewe. The duration of estrus in lambs and in the first estrus of the season is shorter than that of the normal estrus.

27.5.2.4.2 Ram Effect

Most sheep breeding is still done naturally, with the rams pastured with the ewes all year or introduced in the late summer. The introduction of a ram, when the ewes have not been with one tends to synchronize estrus in a high proportion of the ewes 15-17 days later the process is mediated through LH, a pulse of which is released within minutes of exposure to a ram. If contact with the male is maintained, a preovulatory surge of LH at around 36 h occurs, accompanied by a rise in FSH. Ovulation occurs, but no estrous behavior (silent heat). Sexual behavior with subsequent ovulation appears 18 or 25 days after the intro­duction of the ram. The ram effect occurs only in sexually experienced ewes.

27.5.2.4.3 Courtship Behavior

The estrous ewe follows or seeks out a ram, may circle, sniff the male’s body and genitals and then thrust her head against his flanks, fans, or wags her tail, and stands to be mounted. An estrous ewe calls frequently with nonspecific bleats and becomes more active. Standing occurs when the female is receptive, and the ram investigates and nudges her. Like cows, most ewes are in standing estrus at night. Perception via the vomeronasal organ is important. Ewes do differ in their sexual attractiveness to rams and are stable from estrus to estrus. Wooly ewes are preferred to short ones. Further, the attractiveness of estrous ewes for rams depends, in part on the bacterial flora present in their vagina. The bacteria may react with pheromones or themselves contributing to attractivity.

A ritualized kicking with a foreleg is performed as the ram orients himself behind the ewe; the leg is raised and lowered in a stiff-legged striking manner. Tongue flicking accompanies the nudging. The head is tilted and lowered while the shoulder is brought into contact with or oriented toward the flank of the ewe; simultaneously, the ram utters low-pitched vocalizations or gargling or courting grunts. Several abortive mounts may be made with pelvic thrusting but without intromission. When the tip of the glans penis contacts the vulvar mucosa, a strong pelvic thrust accomplishes intromission and ejaculation occurs immediately.

27.5.2.5 Goats

27.5.2.5.1 Doe

Goats, like sheep, are short-day breeders. Does in estrus show frequent wagging of the tail, vocalization, urination, and mounting of other females. This is probably proceptive behavior in that it does not occur as frequently if males are in the same enclosure. Does are in estrus for 39 h once every 20-21 days. There is a male effect in goats similar to that in sheep. Contact with bucks induces an immediate increase of luteinizing hormone followed by ovulation in anestrous goats. In contrast to ewes, a large proportion of does exhibits sexual behavior with the first ovulation.

27.5.2.5.2 Buck

In general, billy goat or buck behavior is similar to ram behavior, kicking at the doe with the front legs and stretching his neck toward her and emitting the gobbling vocalization. The buck holds his tail straight up during courtship. A com­ponent of the mating sequence unique to the goat is the urination by bucks on their own forelegs and beards during courtship. This behavior is termed enurination. Although various functions have been attributed to enurination, includ­ing increasing the intensity of the buck’s odor and advertising his nutritional fitness, it occurs most frequently in a situation of sexual frustration when the buck is restrained from mating. Observation of does mounting one another or copulation by a male and female stimulates ejaculation sooner and with increased frequency.

Silent heat is one of the notable clinical problems in does.

27.5.2.6 Horses

27.5.2.6.1 Mare

27.5.2.6.1.1 EstrousCycle

Mares are long-day breeders and cycle in the spring. Foaling occurs mostly in the late winter and early spring. Mares show individual preferences for stallions, and the preferences are influenced by the stallion’s vocal behavior. The more the stallion neighs, the more likely the mare is to approach him.

The commonly observed problems in mares include anes­trus, split, and prolonged estrus.

27.5.2.6.2 Stallion

Stallions exhibit libido throughout the year but show peak sexual behavior in the spring season. Seasonal changes are also seen in sperm number and testosterone levels. Stallions with a harem have a higher level of testosterone than those in a bachelor herd, which in turn have a higher level than stalled stallions.

27.5.2.6.3 Courtship Behavior in Horses

Courtship behavior will vary with the management practices involved. Driving, herding, or snaking with a distinctive head-down position is behavior usually elicited by the pres­ence of another stallion. An adequate period of sexual fore­play is essential and males may tend to a female for several days before she is fully receptive. Nipping and nuzzling begin at the mare’s head and proceed gradually along the body of the mare to the perineal area. During this testing phase, he exhibits the flehmen response. As sexual excite­ment increases, the male calls with neighs and roars. Full erection usually develops over several minutes in the mature stallion. Several mounts are usually made before intromission and ejaculation. During copulation, the stallion rests his sternum on the mare’s croup and may reach forward to bite her neck. Ejaculation occurs around 15 s after intromission and after approximately seven thrusts, and intromission lasts less than 45 s.

Some common problems of sexual behavior in stallions are lack of sexual interest towards receptive mares, self­mutilation, lack of ejaculation, masturbation, and tendency to injure handlers or mares.

27.5.2.7 Pigs

27.5.2.7.1 Sow

Sows are nonseasonal breeders. After regular cycling commences, the sow will cycle every 18-24 days (average— 21 days) until bred. Puberty occurs at 5-8 months. The presence of a boar leads to the occurrence of estrus at an earlier age.

27.5.2.7.1.1 Estrous Cycle

Sow shows an increase in activity nearing the estrus. Urina­tion is frequent, as is calling to the male. The call is a soft, rhythmic grunt. An estrous female approaches the boar and sniffs him around the head and genitals. Estrous sows attempt to mount other estrous females, but subordinate sows rarely mount dominant ones.

27.5.2.7.2 Boar

Pigs are unusual in that defeminization occurs well after birth and is under the control of estrogenic metabolites that act as late as 3 months postnatally.

27.5.2.7.3 Courtship Behavior in Pigs

After contact with an estrous female has been made, the boar will pursue the female, attempting to nose her sides, flanks, and vulva. Unique to the pig is the boar’s “courting song,” which is used during this phase of courtship. This is a series of soft, guttural grunts, about 6-8/s. Tactile stimulation of the female continues and increases in intensity as the boar’s sexual excitement increases. The boar usually emits urine rhythmically; Pheromones in the urine may further increase the female’s willingness to stand. Several mounting attempts might be made until the female becomes immobile, after which mounting and intromission follow rapidly. Ejaculation occurs within 3-20 min, with an average of 4-5 min.

27.5.2.8 Dogs

27.5.2.8.1 Bitch

27.5.2.8.1.1 Estrous Cycle

The domestic dog, unlike most of its canid relatives, is a nonseasonal breeder. The length of each estrous cycle is extremely variable from individual to individual and some­times from one heat to the next in the same bitch. Usually, one to four cycles yearly may be seen, with two being the most usual.

27.5.2.8.1.2 Courtship Behavior in Bitches

The hormonal levels and sexual behavior is very highly correlated in dogs. The first proestrus and estrus of a bitch’s life are shorter than subsequent ones and the levels of LH and estradiol are lower. She is less attractive to the males and less proceptive. Courtship behavior is marked by play behavior in the proestrous part of the cycle, but this play behavior decreases during estrus. Urination becomes more frequent as estrus approaches and the posture used will frequently be the squat raise. The attraction of males and proceptivity appear in proestrus, but receptivity occurs later, during estrus. During estrus, she stands more quietly to allow male investi­gation and eventually intromission toward the end of estrus. When the male touches her vulva, she will flex her body laterally; while he is thrusting, she will move her perineum so as to increase the probability of intromission. After the lock or the copulatory tie has been established, she may roll or twist and turn.

27.5.2.8.2 Male Dogs

Sexual behaviors may appear in 5-week-old male pups, and mounting behavior becomes an important part of the male’s social repertoire as it matures. As with many other mammals, mounting is used as a sign of dominance; a submissive animal will stand for a more dominant male.

27.5.2.8.2.1 Courtship Behavior in Dogs

Male dogs are attracted to estrous bitches. The urine of the estrous bitch appears to be more attractive to the dog than vaginal secretions, but a component of the vaginal secretions, methyl p-hydroxybenzoate, induces male sexual behavior and is considered as pheromone acting as a “releaser” of sexual behavior in the male.

Males may show extreme interest or indifference to females, although mating may occur successfully in either case. Play behavior may be marked or absent. The male sniffs the female’s head and vulva; he may lick her ears. While canids do not show the classic flehmen response of ungulates, it is possible that the “tonguing” response seen during this olfactory investigation accomplishes the transport of pheromones to the vomeronasal organ in a manner similar to that postulated for ungulates. The male mounts in response to female immobility, thrusts his pelvis and when intromission has been achieved, the rate of thrusting increases. Engorgement of the bulbus gland and contraction of the vaginal muscles following intromission result in the copulatory lock or tie, a phenomenon most closely associated with canids but not restricted to them. The male will usually dismount and turn around so that male and female are facing opposite directions while ejaculation occurs. The lock may last 10-30 min, after which the bulbus decreases in size and the pair separates.

Impotence and lack of socialization are common sexual behavioral problems in dogs.

27.5.2.9 Cats

27.5.2.9.1 Queen

27.5.2.9.1.1 Estrous Cycle

The queen is seasonally polyestrous, and most cats exhibit cycle at least twice yearly if not bred. If unbred, the cat will cycle every 3 weeks for several months. Actual estrus lasts

9- 10 days without copulation and around 4 days if the cat is bred. Most felids, including domestic cats, are induced ovulators, and thus breeding may be accomplished whenever the female shows receptivity.

27.5.2.9.1.2 Courtship Behavior in Female Cats

Anestrous females will call and purr, will be restless, and show increased general motor activity. If she is a house cat she may run from one room to the next, stopping to call at each door or window. She may be very affectionate towards the owners. Urination occurs frequently, and she may spray. She rubs her head and flanks on furniture; glands in these areas may produce pheromones indicative of estrous. She crouches, elevates her perineal region, and treads with her back legs. During proestrus, she will roll and solicit the male’s attention but act aggressively if he mounts. This may be termed postural acceptance and affective rejection. When fully receptive, she becomes immobile and stands crouched in lumbar lordosis with her tail deviated to one side. An estrous female may show a darting behavior in the presence of several tomcats. She will repeatedly run a short distance from the toms, and this may be her means of assessing the relative strength of the males as they chase her and try to displace one another.

27.5.2.9.2 Tom

27.5.2.9.2.1 Courtship Behavior in Male Cats

The male probably locates an estrous female via olfactory cues deposited as pheromones in the urine and by sebaceous gland secretions. A male placed with a female for mating will spend some time investigating and marking the area with urine and anal gland secretions before mating. The cat shows a flehmen response, or gape, similar to that of ungulates. He calls to the female, circles her, and sniffs her genitalia. A nonreceptive female will actively, even violently, rebuff a male. When a female is receptive, the male approaches her from the side and behind and then mounts with the front legs. Intromission follows a forward stepping with arched back and pelvic thrusts. Ejaculation occurs seconds after intromission, and intromission usually lasts less than 10 s. The penis is covered with numerous small spines that apparently cause an intense stimulation as evidenced by the loud copulatory cry of the female with intromission. Copulation may occur every 10-15 min for sev­eral hours.

27.5.3 Maternal Behavior: Formation of Bond Between Mother and Fetus, Concept of Critical Period, Vocalization

Maternal behavior is influenced by hereditary, experiential, neural, and hormonal factors. The combination of the proper hormonal milieu and the stimulus for maternal behavior, the neonate, plus prior experience of being a mother can elicit and influence maternal behavior. The stimulation of maternal behavior appears to be under both hormonal and neural control. E.g., in ewes, estrogen rises and progesterone falls at parturition and hormonal priming by estrogen and proges­terone, plus vaginocervical stimulation, is necessary in order to reduce aggression towards, or withdrawal from, alien lambs by ewes. Experience is also necessary for full expres­sion of maternal behavior because only multiparous ewes would show positive maternal behavior. The fact that primip- arous ewes routinely reject their lambs if they have been delivered by cesarean section also indicates the importance of neural stimulation by the passage of the lamb through the vaginal canal. The fact that multiparous ewes will readily accept their lambs even if they have been delivered by cesar­ean section indicates the importance of prior experience in ovine maternal behavior. The presence of neonates can induce maternal behavior in virgin females and also in males. This phenomenon is called concaveation.

27.5.3.1 Cattle

27.5.3.1.1 Parturition

A greater proportion of births will occur during the day if the cows are fed late at night. Signs of nearing parturition are relaxation of the sacrosciatic ligament, distention of the udder and teats, slackening of the tissue of the perineum and vulva, and mucous discharge from the vulva. Cows will alternate standing and lying much more frequently in the hours before parturition. Arching of the back and tail elevation occurs for 1-3 h before the chorioallantoic membrane ruptures. When the membrane ruptures, the cow often licks the fluid. A periparturient cow will sniff and lick other calves, especially if she is within 24 h of parturition and the other calf has just been delivered, whereas, after parturition, all her activities are directed toward her own calf. Licking of alien calves does neither cause rejection of the cow’s own calf nor does suck­ling of an alien mother cause a calf to fail to suckle its own dam. Other cows will sometimes push or butt a newborn calf. Placentophagia occurs in approximately 82% of cattle.

27.5.3.1.2 Postpartum Behavior

Contact between the cow and her calf for as brief a period as 5 min postpartum results in the formation of a strong, specific maternal bond. Cows groom their calves during the early postpartum period, especially on the back and abdomen. Licking the calf occupies up to half the cow’s time during the first hour postpartum; heifers lick less. Licking not only dries and stimulates the calf but also results in analgesia in the cow.

As soon as the calf’s shoulders are free of the mother’s vulva, the newborn calf begins to shake its head, snuffle, and sneeze. Some calves remain motionless for up to 30 min after birth, but within an hour most calves can stand. For the first time, the calf may take 30 min to an hour to locate the teats, but it will be able to locate it more quickly in subsequent nursings. Sucking problems and Cross-fostering are common abnormal maternal behavior.

27.5.3.2 Sheep

27.5.3.2.1 Parturition

Maternal behavior in sheep has an important clinical rele­vance, as most lamb mortality occurs within the first week of life in range-reared sheep. Lambs may be born at any time of the day or night, with peak frequencies being noted between 9 and 12 in the morning and between 3 and 6 in the evening. A few days before parturition, the ewe withdraws from the flock and seeks shelter. Shelter seeking by the ewe improves the environment into which the lamb is born so that its chances of survival are better. Nearing parturition, the ewe will withdraw and seek a corner in the pen. She will show restlessness, circle, vocalize, rub her head on her flanks, lick herself, and paw at her bedding 60-90 min before parturition. Grazing and ruminating cease. The interval between the onset of labor and the appearance of the lamb is usually 30-60 min but may vary. Even before parturition, nearly 20% of ewes show maternal behavior toward other lambs. This prepartum maternal behavior results in lamb stealing.

27.5.3.2.2 Postpartum Behavior

When the lamb is born, the ewe begins to lick it during most of the first hour after parturition. Primiparous ewes bleat more. Licking of the lamb is very important especially in cold or windy weather because it serves to dry the neonate and it additionally serves to stimulate the lamb. While the lamb is recumbent, the ewe licks its head, even restraining the lamb with a front leg to prevent it from standing. Licking of the perineal area stimulates the lamb to rise. Usually within the first 30 min, the lamb tries to stand and the ewe continues to lick the lamb mostly on the hindquarters. If the ewe stops licking, the lamb gives distress calls. Finally, licking of the lamb by the ewe establishes the maternal-offspring bond, which enables to identify her lamb by smell and taste. Usu­ally, the fetal membranes are licked off the lamb and ingested, but the placenta is not eaten. The lamb raises and shakes its head and bleats. Although lambs are able to stand within 30 min or an hour of birth, it may take 2-3 h before they find the udder.

The “critical period” during which an ewe will accept a lamb is the first several hours after parturition. Normally, a ewe will stay within two meters of her lamb for most of the first day. If a ewe’s lamb is removed immediately after birth, before she has licked it, the ewe will accept any lamb presented to her. After the ewe has spent 30 min to 2 h with a lamb, her own or a substitute, she will not accept any other lamb. Mutual recognition by the ewe and lamb depends on olfaction, audition, and visual cues. Common maternal behavioral problems include Cross-fostering, Mismothering, and poor mothering.

27.5.3.3 Goats

As parturition approaches, does, especially multiparous ones, leave the herd and seek a sheltered place, almost always near a vertical object. The does will defend this area both before and for the first day after the kid is born. A doe is usually very agitated vocalizing, urinating, and moving, but this response disappears just before parturition. As parturition approaches, they grunt, paw the ground, kick, and lick their backs. Does usually lie down for kidding. After parturition, the kid is sniffed and immediately licked; the head is the primary target. The licking continues for 2-4 h.

Goat kids should stand within 20 min and suckle within an hour. Within 4 h after the kid’s birth, the doe can recognize it by sight and sound. Two-day-old kids can identify their mothers visually. Olfaction seems to be important for selec­tive maternal behavior. Kid rejection is one of the most common maternal behavioral problems in goats.

27.5.3.4 Horses

27.5.3.4.1 Parturition

The onset of parturition in mares is heralded by waxing of the udder, but the length of time between the appearance of udder waxing and the appearance of the foal might be quite variable, up to 21 days. Body temperature is lower on the day prior to parturition. The mare will walk more and stand less on the evening of parturition. In the first stage of labor, which lasts for about 4 h, the mare is restless and will crouch, straddle, and urinate. The smell of fetal fluids is attractive to parturient mares. The mare will exhibit the flehmen response in response to the amniotic fluid that is expelled. Sweat will appear on her elbows and flanks. During the second stage of labor, the mare will lie in lateral recumbency. This second stage is very violent and very short in horses, lasting less than a half hour.

27.5.3.4.2 Post-Parturient Behavior

Ordinarily, the foal is delivered in such a way that the mare needs only turn her head to meet her foal muzzle to muzzle. Licking, as well as sniffing, is concentrated first on the head of the foal and later on the hindquarters, particularly the perianal area.

Standing and suckling occur within 1-2 after birth for pony foals. Foals suckle four times per hour at 1 week of age and gradually decrease the frequency to once per hour by 5 months. Mares are very protective of their foals and are usually very aggressive towards other horses and sometimes towards people for the first day or two after foaling. The first hour is probably critical for the mare to learn to recognize her foal selectively. The foal appears to take much longer, per­haps as long as a week, to recognize the mare. The neighs (or whinnies) of the separated mare and foal are impressive, and horses make use of these calls to locate one another. Mismothering can occur in equids, although it is much rarer than in sheep.

27.5.3.5 Pigs

27.5.3.5.1 Parturition

Most of the farrowing take place in the afternoon or night. After labor begins, most sows lie in lateral recumbency. The sow will swish her tail violently as abdominal straining takes place. Parturition usually takes 3-4 h but varies considerably with litter size and the condition of the gilt.

27.5.3.5.2 Behavior of the Sow

Maternal behavior in sows is exhibited as calmness and care when lying down to avoid piglets and remaining in the nursing posture following milk ejection and protectiveness in response to piglet squeals and human approach to piglets. When not confined, the sow will eat the placenta. It may be recycling of nutrients or a form of defense against predators by removing odors or as a means of inducing analgesia. Sows do not often lick their newborns.

27.5.3.5.3 Behavior of the Neonatal Piglet

Piglets make a most startling transition from fetal to indepen­dent existence. They may be apnoeic for 5-10 s after birth. Then, they give a few gasps before beginning to breathe regularly. Their eyes and ears are open, and they are able to walk immediately, although their gait is staggering for the first few hours. Most piglets are nursed within 30 min of birth. Piglets are attracted to soft, warm surfaces, pig vocalizations, and the sow’s odors and they move in the direction of the sow’s hair growth. The firstborn piglet appears to use thermal, tactile, and olfactory cues to find the udder, whereas the later-born probably respond to the suck­ling sounds of their older littermates and walk straight to it because social facilitation is strong in pigs at birth.

27.5.3.5.4 Mutual Recognition

Sows and piglets apparently use olfaction to identify one another but need more than a day and possibly as long as a week to learn. The piglets can identify their dam’s feces, milk, and urine odors, as well as their vocalizations. Sows respond to playback of piglet separation calls by vocalizing, but cannot discriminate their own from other piglets on the basis of their voices. However, she learns to identify her own piglets by the time they are a week old on the basis of olfaction. Piglets can easily be fostered by another sow when they are less than a day old. After that, the fostered piglets walk around, and are reluctant to suckle, as they have already formed a bond with their dam.

Cannibalism occasionally occurs in sows, especially in nervous primiparous gilts and sows suffering from mastitis will refuse to nurse.

27.5.3.6 Dogs

27.5.3.6.1 Parturition

During late pregnancy, the activity of the animal decreases and its appetite increases. The animal will be anorectic as abdominal pressure increases as it nears whelping and takes only small quantities of feed. The animal will be usually restless, shows nesting behavior, and grunts each time it sits, especially if it is carrying a large litter. At parturition, the bitch licks itself, any soiled bedding, and, almost inciden­tally, the puppy. The pup’s head, umbilicus, and perineum are the area’s most frequently licked. A pup is delivered every 30-60 min on average. The bitch cuts the umbilical cord with her molars and then licks the stump. The placenta is expelled 5-15 min after each pup is born. If the placenta has not been delivered, it may extract it by pulling on the cord. The placenta is usually eaten by the bitch. As the labor progresses the animal usually becomes silent and less active unless dystocia occurs. Labor can be interrupted if the bitch is disturbed. Depending upon the temperament of the dog, 15 min to an hour is required for normal parturition to proceed. Amniotic fluid is very important because if pups have been cleaned the bitch will not accept them.

27.5.3.6.2 Postpartum Behavior

The puppies move forward by paddling and after they encounter the mother ’s body, they nuzzle through the fur, usually attempting to burrow beneath her. Puppies have two vocal signals, the whine and the grunt. The whine is emitted whenever the puppy is cold, hungry, or separated from its litter or mother. The grunt is apparently a pleasure communi­cation that occurs when sought-after warmth or reunion occurs.

Bitches lick their puppies a great deal. The licking arouses the pups to eat, and, when it is directed at the anogenital area, it stimulates urination and defecation that would otherwise not occur spontaneously. The bitch keeps the nest area clean by consuming the urine and feces of her puppies for at least the first 3 weeks of their lives. Licking helps in retrieval which is unique to dogs. Bitches usually do not carry their puppies. Instead, they lead them back to the nest by licking the pup’s head. The common problems encountered are pseudopregnancy, maternal rejection, and repeatedly moving the pups.

27.5.3.7 Cats

27.5.3.7.1 Parturition

Gestation period in cats is 63-66 days. Births have a seasonal distribution, with the greatest number of litters being born in the summer and the least in autumn and early winter.

Most cats will choose a cave-like place, such as a closet or a linen cabinet. Parturition in the cat is characterized by a great amount of licking by the queen: self-licking mostly directed at the belly and genital area, licking of the fetal fluids from her body or the floor, and licking the kittens. The queen is typically very restless and other behaviors include frequent lying down, sitting up, squatting, bracing lordosis, licking of the vulva, circling, and walking around the cage.

The queen will eat the placenta as it is delivered. In the process of eating the placenta, the queen stretches the umbil­ical cord so that little bleeding occurs when the cord is severed. Most kittens are suckling within an hour or two of birth.

27.5.3.7.2 Postpartum Behavior

Even before the opening of their eyes, kittens can return to their nest from several feet away by using olfactory cues. Queens retrieve their kittens in response to auditory, rather than visual signals. The more the kitten vocalizes, the more apt the queen is to retrieve it. The queen usually picks the kitten up by the scruff of the neck, though occasionally it grasps the skin at the back of the head or even the kitten’s whole head.

Grooming plays an important part in feline maternal behavior as in most other species. Queens lick their kittens frequently and in particular lick the perineum to stimulate urination and defecation for the first 2-3 weeks of life. In common with most carnivores, female cats ingest the kittens “urine and feces” for several weeks postpartum thereby keeping the nest clean. Cats will accept alien kittens that are not too much older than their own at the time of parturition. Infanticide, mismothering, and overgrooming are the com­mon problems observed.

27.5.3.7.3 Parental Care and Its Influence on Behavior Parental influence and parental contacts vary among different species. For example, marsupial infants are born after a very brief gestation period and they are fused to the mother’s nipples and are no more than externalized fetuses. Their sensory capacities are very much limited at this stage and mother-infant interactions is only through nursing. But even when they become independent, they are constantly with the mother, carried in the pouch thus representing a far higher degree of contact than the part-time attention received by baby rats and mice.

In contrast, the mother rabbits (Oryctolagus) visit their litter only once a day initially. and the milk is pumped into the babies. The young ones are briefly groomed and then the nest is covered and left; the mother herself has another nest chamber elsewhere. Probably this behavior helps to minimize the chances of predators finding the nest by picking up the mother’s scent.

A similar explanation is suggested for the yet more extreme maternal behavior of tree shrews (Tupaidae), a rather distinct group of small insectivore-like creatures that have a distant linkage to the primates. They build two separate nests, in one of which the adults sleep while the female produces the litter in the other. The young are visited briefly every second day; they suckle rapidly and become distended with milk. The mother then leaves them with the very minimum of grooming and typically does not return for 48 h. The young are capable of grooming and licking themselves from the day of birth. They stay alone in the nest until they emerge at 33 days of age.

The primates, have some of the longest and closest associations between parents and offspring, lasting for 6 or 7 years and beyond in the great apes. The relationship between a primate mother and its offspring is absolutely crucial for almost every aspect of behavioral development. Separation, particularly in the early months, can have very severe effects on infants.

27.5.4 Milking Behavior and Factors Influencing Milking Behavior

Milk let-down is a neuroendocrine reflex initiated by oxyto­cin release following stimulation of the teat by a suckling calf. However, the cows with calves soon start to release oxytocin when other stimuli from the calf are detected, and cows milked using a milking machine may respond to other cues, such as the sounds, in a similar manner. Milk let-down by dairy cows in response to the typical sounds of a milking parlor is the best example of classical conditioning on farms. Milk let-down is a conditioned response to the conditioned stimulus of clanking noises, made by the washing of milking vessels, etc., in the milking parlor.

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Rhesus monkeys are more restrictive, rarely allowing their babies to leave their side for several weeks and control outside contact for a much longer period. How­ever, the Barbary macaque (Macacasylvanus) of North

Africa which is a close relative to the rhesus monkeys behaves quite differently. Mothers allow other individuals, males as well as females, to hold their babies even within a few days of birth. In all primates, it is commonly observed that newborns and young generally are objects of intense interest to other members of a group, particularly young females. “Aunting” behavior has frequently been reported from primates and is also an important part of parental behavior within the matriarchal groups of African elephants. They too have a long infancy and a rich social environment. Elephant calves do their best if they grow up in a group where they have not just their mother ’s attention but of other females and juveniles as well.

There is a breed variation among cattle with respect to how readily such conditioning can occur, for example, breeds such as the Salers in France do not readily let-down milk to stimuli other than those emanating from real calves as com­pared to Friesians or Holsteins. Farmers need to be aware of the fact that milk let-down in a parlor is a conditioned response and that such learning depends upon adequate experience and training.

27.5.4.1 Factors Influencing Milking Behavior

• Animals should not be handled roughly or beaten while leading to the milking parlor. They should not be coerced into the milking area.

• A frightened or stressed animal may show inhibition in the conditioned milk let-down. Experiments have shown bet­ter milk let-down in response to music played in the milking area.

• Strangers should not be allowed in the milking area during milking.

• Presence of disturbing stimuli in the milking parlor may prevent the young animal from learning and the older animal that is conditioned may be inhibited from exhibiting the response.

• Veterinary operation involving discomfort for the cow should be avoided out in the milking parlor and carried out in a separate facility.

• Providing concentrates during milking may strengthen the conditioned milk let-down.

• Barking dogs may be avoided in the milking area.

Nervous cows that are difficult to get into the bail or those not settling down for easy milking will not be suitable as good milking cows. These may be replaced with cows which have a quiet and relaxed temperament.

27.5.5 Social/Gregarious Behavior

In wild, animals often tend to prefer living in social groups as it provides protection against predator attacks, enhance hunt­ing prospective, mating opportunities, helps in learning, finding feed, grasslands, and water resources. Animals in groups exhibit various behaviors such as dominance hierar­chy, peck order, teaching, cooperation, and playing.

27.5.5.1 Dominance Hierarchy

It is a type of social hierarchy that arises when members of a social group interact with one another, to create a ranking system. In social living animals especially in wild, members compete for limited feed resources and mating opportunities. Rather than fighting they confront; a relative rank order is established between the members of the same sex in a spe­cies. This order is thus created based on repetitive social interactions and is subject to change whenever the dominant animal is challenged by his/her subordinate. Hence, these hierarchies are not fixed and depend on a number of changing factors such as age, sex, body size, intelligence, and aggressiveness.

Several brain areas are linked to hierarchical behavior in animals. One of the important areas is prefrontal cortex, a region involved in decision-making and social behavior. High-ranking macaques have a larger rostral prefrontal cortex in larger social groups. Amygdala, thalamus, and dorsal raphae nucleus are also associated with social ranking behavior.

27.5.5.2 Peck Order in Chickens

The social rank order established in chicken is referred as peck order. The peck order influences the feeding, drinking, egg laying, mating, and crowding activities. The stronger members of a flock are at the top of peck order while weaker or submissive birds are placed lower in the order. Hens high in the pecking order chase out other hens out of nest boxes they favor. There is a reduction in aggression and a lower incidence of conflicts when a peck order is established.

27.5.5.3 Teaching in Animals

Teaching is a behavior often seen in animals and it helps in imparting some new information to offsprings/individuals faster than they would otherwise receive it. Mother cheetah captures live prey and allows its young to interact with this prey, making sure that the prey does not escape along the way.

Another example can be observed wherein the meerkats young pups begin following groups of foragers, and the pups are assisted in their own foraging attempts by older groupmates called “helpers.” Helpers will often incapacitate scorpions by removing their stingers and presenting them to pups as food. Very young pups were either fed dead or incapacitated scorpions, but as the pups got older, the helpers presented them more and more often with live scorpions.

Similarly, the key element of elephant society is the matri­archal group led by the oldest and largest female with her daughters and their offsprings. A baby born into a group is known to all others, it is nourished and closely protected by its mother and other relatives for several years. Slowly it acquires the adult feeding behavior, learning how to select food and migrate according to the seasonal changes of vege­tation and water supply.

27.5.5.4 Cooperation

Cooperation refers to an outcome in which two or more interacting individuals receive a net benefit from their joint actions, despite the costs they may have to pay for undertak­ing such actions. For example, jointly hunting prey may provide each of the two hunters with food, even though there are costs (possible injury, energy expended) associated with hunting. In addition to looking at outcomes (i.e., suc­cessfully capturing prey), it is also important to examine cooperation in terms of individual action.

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An interesting case of cooperation is observed in the Rodriquez fruit bat (Pteropus rodricensis). In this bat, during the birthing process, unrelated female helper continues providing assistance by grasping the wings of the pregnant female, providing both protection and warmth, and subsequently cleaning (licking) newborn pups upon their emergence. Once pups are born, helpers guide the newborns into a suckling position, where they can obtain milk from their mother. These bat “midwives” cooperate with pregnant mothers dur­ing virtually every stage of the birthing process. Assis­tance during birth has been recorded in marmosets, Indian elephants, African hunting dogs, raccoon dogs, and bottle-nosed dolphins.

27.5.5.5 Social Grooming

Many animals groom, or clean, themselves in an attempt to remove external parasites. This sort of grooming might involve scratching an area of skin or licking it. Social grooming, or allogrooming, in which one individual grooms another, is one of the most obvious and frequently observed cooperative behaviors. While social grooming often serves the function of removing parasites from the body of a partner, it may also have many other functions as well. Social grooming has been considered a large part of the glue that holds primate troops together. One benefit linked to social grooming in primate groups is “tension reduction.” One way in which primates of a group reduce the chances of escalating violence is through social grooming, which has the effect of lowering the level of tension between putative combatants. From a proximate perspective, one means by which social grooming lowers tension levels is by increasing the circulating levels of hormones such as endorphins and opioids. This sort of tension reduction through social grooming may play a role in nonprimate species as well. Primates also appear to exchange social grooming for access to scarce resources such as water or food, entrance into new groups, aid in chasing potential predators away, and future association with individuals who possess “special skills” that they themselves do not.

Other examples of specialized behavioral learning include food-storing behavior, i.e., when the chance arises, a number of bird and mammal species collect more food than they can eat immediately and hide it. In this way, they can take advantage of a rich food source while it is there and reduce the degree to which they have to share it with others. Some animals make a single larder to which they return regularly, interesting some species hide food items singly or in caches dispersed around their territory.

27.5.5.6 Play

Animals play with each other and with objects spending long periods. Although the adults may continue to play intermit­tently, it tends to fade out with age. It is commonly suggested that play has a role in the development of adult behavior. Play is an energy-consuming activity and 5-20% of the “surplus” energy available is expended while playing. The cost is not just in energy; young desert bighorn sheep (Ovis canadensis) cavorting in play through their harsh environment are often pierced by cactus thorns; young gelada baboons (Theropithecus gelada) playing on the steep cliffs of Ethio­pian gorges have been noticed to fall 5-10 m and limp painfully away. Another significant cost arises because vigi­lance is likely to be reduced when young animals play, making it potentially dangerous.

The play behavior is very conspicuous among the ungulates, carnivores, whales, elephants, and primates. Each group has its own typical play repertoire; ungulates butt each other, mock fights and chases are common among primates, while carnivores show elaborate stalking, leaping, and prey-catching behavior. The mock fights among kittens or the amazing high-speed chases and wrestling bouts of young monkeys play a vital part in the development of skills which are going to become vital for survival during adult life.

Apart from the obvious physical skills, numerous other functions have been suggested for play such as gaining knowledge about potential prey species, gaining knowledge of the social group and one’s position within it, exploration of the environment, and so on. It prepares animals for the unexpected. Mothers often take part in play sessions and their influence can be detected indirectly as well. Play builds up the flexibility of responsiveness and hence improves sur­vival. Play in young mammals takes place at a time when crucial maturation events are occurring in the brain and muscles which lead to the differentiation of synapses in the cerebellum and the control of complex movements.

With some members of the cat family, there is a very conspicuous type of play that seems particularly directed towards the teaching of prey-catching. Domestic cats bring home live prey which they “play” with, letting them loose only to pounce on them again, often repeatedly. Cheetah mothers may bring back a crippled gazelle fawn to her young ones.

27.5.6 Agonistic (Combat or Aggressive) Behavior

In animals, aggression is a threat or harmful action directed against another individual and includes snarling, growling, and biting. Aggressive behavior is usually displayed to threaten or attack to resolve competitive disputes over limited resources (feed, territory) or to increase their reproductive potential, or to escape from threatening situations. However, aggression can have multiple motivations, especially in pet animals.

Aggression is a normal communication modality and is not necessarily pathological or abnormal behavior. Normal aggression is demonstrated in situations that warrant aggres­sion. The intensity of the aggression by the animal is modified, depending on the situation and the relative level of the threat. For example, a normal aggressive display would be if a stranger broke into the home unannounced and was bitten.

27.5.6.1 Conflict-Induced Aggression

This aggression typically occurs when the pet seeks out the pet owner, perhaps even solicits attention, then shows con­flict behaviors (freezes, shifts eyes) and aggresses. The pet could leave the situation but chooses not to because of conflicting motivations (desire to be with the pet owner and uncertainty of what will happen next.)

27.5.6.2 Possessive Aggression

Possessive aggression is described as guarding of an item (food) from people.

27.5.6.3 Alliance-Induced Aggression

Alliance-induced aggression refers to aggression between dogs in the same household that fight only in the proximity of the pet owner. Usually, the more fearful and most owner­dependent dog becomes anxious when the other dog approaches the owner and consequently the less confident dog aggresses.

27.5.6.4 Idiopathic Aggression

Idiopathic aggression refers to an unexplained and intense aggression in which no identifiable aggression-provoking stimuli can be determined on a complete medical investigation.

27.5.6.5 Maternal/Hormonal-Induced Aggression

Maternal aggression occurs when a bitch defends her puppies from people or other animals. It is considered to be normal behavior to a certain degree.

27.5.6.6 Play-Induced Aggression

Canine mouthing and play biting are considered normal forms of interactions and communication for dogs but can become problematic when directed toward humans. Although not considered as a true aggression, over-exuberant play can be damaging to the human-animal bond.

27.5.6.7 Redirected Aggression

Redirected aggression is incidental to another form of aggres­sion or emotional arousal. It occurs when a dog or cat cannot reach the target of its aggression and out of frustration changes its focus of aggression to an object, person, or other animal that is not the stimulus for the aggressive arousal.

27.5.6.8 Territorial Aggression

Dogs are genetically programmed to be territorial. Territorial aggression refers to the aggression directed towards people or animals that are not part of the dog’s immediate social group or are unfamiliar to them when they enter the dog’s perceived territory. The territory may be a fixed location (i.e., house, room, yard, and bed) or a mobile territory around the dog (i.e., leash, proximity to owner). Territorial aggression is influenced by breed, age, sex, and socialization experience.

27.5.7 Communicating Behavior and Modes of Communication in Different Species

Animals communicate not only by auditory signals but also by visual and olfactory signals. Communication in animals depends on their capacity to perceive messages. Many of the stimuli animals encounter come from other animals and these form the basis of communication. Animals communicate with each other using signals, which often closely match the sensitivities of the sense organs of the animal receiving them.

27.5.7.1 Visual Signals

Every animal communicates with one another, with animals of other species through various visual cues. The body pos­ture movements and overall behavior differ among species.

27.5.7.1.1 Dogs

A dog’s emotional state can be identified by observation of its ears, mouth, facial expression, tail, and overall body position and posture. A calm dog stands with ears and tail hanging down. When it becomes alert, its tail and ears are pointed upward. The dog may point with one front foot and as the dog becomes more aggressive, the hair on the shoulders and the rump rise and the lips are drawn back. The ears remain forward and the tail wags slowly. On increasing aggression, the lips get retracted and the teeth are exposed in a snarl. The dog stands straight. As the dog gets frightened, the ears go back and are flattened against the head and the tail is tucked between the legs.

Dogs greet their owners as they do with their mothers: by licking their faces. Puppies lick their mothers’ faces begging for regurgitated feed.

27.5.7.1.2 Cats

A cat carries its tail high when greeting, investigating, or frustrated. During stalking, the tail is depressed and the tip is wagged. A calm and relaxed cat usually stands with its tail hanging; its ears are usually kept forward.

27.5.7.1.3 Pigs

The tail, particularly in piglets, is a good index of their general well-being in most breeds. Although Vietnamese mini pigs do not curl their tails, a tightly curled tail indicates a healthy pig in most breeds, and a straight one indicates some sort of distress.

27.5.7.1.4 Horses

The horse’s ears are probably the best indicator of its emotions. The alert horse looks directly at the object of interest and holds its ears forward. During aggression, the ears are pointed back, and the flatter the ears are against the head, the more aggressive the horse is.

27.5.7.2 Vocalizations

The animals vocalize with one another to communicate.

27.5.7.2.1 Cattle, Sheep, and Goats

Vocal communication in prey species such as cattle may be the most important in disseminating information about gen­eral safety or danger. It might have been more important for cattle to be alert and ready to flee than to communicate more precise information in their calls. The moo is low-pitched and most common. The other common vocalizations are higher pitched—hoot or roar consisting of repeated brief calls, usu­ally by a distressed cow. A threatening bull gives a roar of high amplitude. A very hungry calf will give a high-intensity “menh” call. Grunting sounds are heard during copulation. Vocal communication in sheep consists of bleating in distress or to initiate contact. Ewes rumble to their newborn lambs and rams make a similar call while courting. The snort is an aggressive communication in sheep.

27.5.7.2.2 Dogs

The common vocal communications of dogs are bark, whine, howl, and growl.

Barking is a territorial call of dogs. It is used to guard a territory and to demarcate its boundaries. Whining is a care­soliciting call of the dog used by puppies to communicate with the mother, who provides warmth and nourishment. Mature dogs whine when they need relief from pain or are in even a mildly frustrating situation, such as when they want to escape outdoors or reach a rabbit for which they are digging. Howling is a canine call that occurs more frequently in wild canids, coyotes, and wolves and in some breeds of dogs, such as huskies, malamutes, and to a lesser extent in hounds. Growling is an aggressive or distance-increasing call in dogs.

27.5.7.2.3 Cats

The common vocalization call include Murmur, Purr, Growl, Squeak, Hiss, Shriek, Spit, Estrus call, and Chatter. Howl and yowl of an aggressive cat, Mowl, or caterwaul, Mew, Moan, and Meow.

27.5.7.2.4 Pigs

Vocal signals are probably the most important means of communication in pigs. Nearly, 20 calls have been identified, Bark, Grunt, and squeal being the common ones.

Bark is given by a startled pig. A shorter grunt is given by an excited or investigating pig. Continuous short grunts are given by a threatening sow and may precede an attack on anyone who disturbs her litter. In a milder form, it can be a greeting. A long grunt (0.4-1.2 s) might be a contact call and is often associated with pleasurable stimuli, especially tactile ones. The squeal is a more intense vocalization indicating arousal and an injured pig screams.

27.5.7.2.5 Horses

Neigh and nicker are common vocal calls which evoke a reply. The neigh (or whinny) is a greeting or separation call that is most often heard when adult horses or a mare and foal are separated. The soft nicker is a care-giving or care­soliciting call given by a mare to her foal upon reunion and probably is mutually recognized. A horse may also nicker to its caretaker and a stallion to a mare in estrus.

27.5.7.3 Olfaction

Olfactory acuity is probably the most important sense in the domestic animals because individual odor recognition and pheromonal release are an important part of their communi­cation and especially during breeding. Olfaction to animals is what writing is to humans, a message that can be transmitted in the absence of the sender. The sender must be present for auditory or visual signals to be sent, but an odor persists for minutes (or days) after the sender has gone. Dogs probably have the greatest olfactory acuity and this macrosmatic species hence is the one most investigated. They can distinguish between the odors of identical twins and detect the odors of fingerprints 6 weeks after the fingerprints were placed on the glass. Bloodhounds are indeed the most sensitive of all dog breeds. Dogs frequently are trained to sniff out drugs and natural gas leaks, and bloodhounds have been used for centuries to track people; apparently, no modern invention is as reliable as the canine olfactory mucosa. A large dog has over 15 times the area of the epithelium of a human and 100 times the density of sense cells per unit area. Pigs and bears use their sense of smell to detect roots and bulbs or burrowing rodents well below ground. Males can also determine whether a female is ready to mate from her scent.

27.5.7.3.1 Olfactory Signals in Cats

Cats use several olfactory signals such as scent marking and rubbing with the secretions such as anal secretion, urine, and facial gland secretion.

Male cats scent marks, that is, spray urine, more than females, but both sexes do it. They spray trees along their most frequently traveled path. Spraying is also done by cats that are subjects of aggression. The smell of tomcat urine is due to the sulfur-containing amino acid, felinine, which is an important olfactory component in territorial spraying. Cats are well known for their fastidious covering of their feces, but in some situations, such as outside their core living area, cats may leave their feces uncovered. Cats probably use fecal and anal sac odor for communication. Cheek rubbing (bunting) behavior may also be a form of olfactory communication in that glandular secretion from the cat’s face is deposited on the object bunted.

27.5.7.3.2 Olfactory Signals in Boars

Boars may use behavioral signs more than pheromones to determine the sexual receptivity of the sow. Boars are the only male ungulates that do not exhibit flehmen. Instead, they gape like a cat does when they encounter sow urine. Females can identify intact males, by the strong boar odor produced by the androgen metabolites present in both the saliva and in preputial secretions.

27.5.7.3.3 Other Signals

Duck-billed platypuses, sharks, and rays have the capacity to detect the electric fields generated by muscular activity in animals such as the crayfish and shrimps on which they feed. Electric fish go one stage further and create their own electri­cal environment by using specially modified muscle tissue which helps in social communication and to find prey.

27.5.8 Behavioral Disorders

27.5.8.1 Dogs and Cats

Failing to respond when called, jumping up on owners or visitors, and running away are minor behavior problems which may strain the owner-dog bond. Tail chasing, light chasing, circling, and digging for imaginary prey are com­pulsive problems. Phobias, especially the fear of storms, are often a serious problem. Destructive chewing behavior caused by oral exploration, escape attempts, or separation anxiety are common complaints reported by the owners. Signs of separation anxiety include destruction at doors, general destruction in the house, house soiling, excessive salivation, distress vocalization, or self-trauma. Other problems include urine marking, hyperexcitability, and acute conflict and stereotypical behaviors.

27.5.8.2 Horses

Stall walking in horses may occur as a stereotypy, a repeti­tive, functionless behavior seen mostly in confinement. Pawing is a response to frustration, a displacement activity that might have originated from the activity of uncovering food buried under snow. Most horses kick the stall walls with their hooves and a few horses may knock their hocks against the wall. Cribbing is an oral behavior wherein the horse grasps a horizontal surface, such as the rim of a bucket or the rail of a fence, with its incisors, flexes its neck, and aspirates air into its pharynx. Some horses may aspirate air without grasping an object. This is called aerophagia or windsucking or pneumovaginitis.

As horses are generally neophobic, that is, they are afraid of new or strange things many of them exhibit failure to load when transported on the ground by trailers. Both innate behaviors and learning contribute to loading problems. Dark interior of the trailer; the hollow sound of hooves on the ramp, and the instability of the ramp and vehicle, and unpleasant experiences with loading or riding in a trailer may exaggerate horse’s innate fears. Hence, a horse that is reluc­tant to enter a trailer may be willing to follow another. Horses are less stressed when traveling with a companion horse.

27.5.9 Training of Dogs and Horses

Specially trained working dogs have a variety of roles in our society, ranging from livestock handling and care in farms, to military and law-enforcement dogs who detect bombs, fugitives, or drugs; to search and rescue dogs who save people trapped by natural disasters such as earthquakes or avalanches. Guide and service dogs contribute invaluable assistance to the visually and hearing impaired and people with limited mobility and in recent years they are being trained as therapy dogs to help autistic children, the aged, and people in depression to overcome their health issues and interact with the society in a better way. Human-animal interactions documented to have positive health effects in the areas of social behavior, interpersonal interactions, and mood; improvement in stress reduction scores, heart rate, and blood pressure; and reduction in fear and anxiety.

Training becomes a necessary component in rearing animals, especially in pets and we often need to train them for basic daily life skills and specific purposes-oriented skills. Being highly intelligent social animals, they have proven themselves capable of learning many extraordinary tasks to assist us. Dogs know when we are paying attention to them and understand our perspective. They can convey informa­tion to people as well as read our signals to them. Dogs can signal the location of a hidden object by using “showing behaviors,” like alternating their gaze from the object to their human companion repeatedly until the target is acquired. Studies report that dogs use multiple facial features of humans such as the eyes, midface, and mouth to determine emotional meanings and respond accordingly. Thus, training the animals involves intelligent interaction between the trainer and the animal demanding a lot of patience and understanding.

27.5.9.1 Training in Dogs

There are two types of associative learning that are particu­larly useful in training and working with dogs: classical conditioning and operant conditioning. Classical condition­ing involves reflex-like, involuntary, or emotional responses, and occurs when a stimulus that was previously meaningless (“neutral”) to the dog takes on the power to elicit such a reflex-like response. Practically, we can use classical condi­tioning to build a positive association between the dog and something or someone (i.e., some stimulus or trigger) in the dog’s environment. For example, if we want the dog to be friendly with children, veterinarians and other dogs, and cats, we can achieve it by some classical conditioning, i.e., we may ensure that whatever the trigger is, it becomes a predictor of good things. So, each time the puppy sees a child or veteri­narian approaching on a walk speaks in a happy voice and offer him a delicious treat essentially, we are trying to teach him that “children/veterinarian mean good things are going to happen.”

Another very important form of associative learning is operant conditioning. Behaviors that are followed by positive outcomes (i.e., are rewarded or reinforced in some way) are more likely to be repeated; behaviors that are followed by negative consequences (i.e., result in some form of punish­ment) are less likely to be repeated. One important aspect of using rewards in dog training is the speed at which the dog is rewarded for successfully performing the behavior. If the reward takes longer than a second or two to reach him, he may not understand exactly why he has received the reward (or, he may have performed a different, less desirable behav­ior in the meantime). Provision of immediate reward and reinforcement to the dog can greatly improve the effective­ness of the training.

Shaping a way to train a new behavior, especially a fairly complex one, is to break down the desired behavior into tiny increments or steps, and reinforce the dog at each incremental step until we get the full behavior from the dog.

Continuous reinforcement means that every time the dog performs the behavior, he earns a reward and is the best schedule to use when teaching a dog a new behavior. Inter­mittent reinforcement, on the other hand, means that the dog does not receive a reward every time he does the behavior, but only periodically, and either on a fixed schedule or on a variable schedule. Intermittent reinforcement should only be used once the dog has successfully learned the behavior, but it is often the best way to maintain the acquired response (i.e., the dog will continue to work in anticipation of the reward, even when the reward does not happen each and every time).

Today, rewards-based training is considered as the science in dog training. Older methods that rely heavily on force and positive punishment are not in vogue. Scientific literature on dog training methods and veterinary medicine highlight and emphasize the dangers of positive punishment (increased fear, high risk of increased aggression, disruption to human-animal bond).

27.5.9.2 Training in Horses

While human-horse interaction has existed for many millennia through hunting, it is only relatively recently that horses have been used for agriculture, transport, war, and, more recently, for sport and leisure. Since the beginning of domestication, various techniques for horse training have been developed and passed on to subsequent generations orally or through the literature. However, all these techniques are constrained by the biology of the horse. When it comes to getting the perfect efficient output from the horses in sport and work, we need to be well acquainted with their behavior. Effective and humane training always takes account of the animal’s ethology, even though the training systems differ.

Horse training differs fundamentally from the reward­based training methods used for marine mammals, exotic carnivores, and most companion animals because it largely relies on negative reinforcement. During their early training, horses learn that the correct responses result in the reduction of pressure from the reins when they stop or slow. Pressure from the rider ’s legs is reduced when the horse moves for­ward. To be effective and humane, the application of pressure must be subtle and its removal immediately once the horse complies.

Training a New Behavior in Horse

When a horse is taught a new behavior, the behavior should be reinforced after every occurrence initially and then the reinforcement can be gradually spaced after occurrence. In this manner, the behavior is more likely to be rapidly learned and efficiently remembered well. For this, the horse must achieve the goal in order for it to be rewarded.

An effective trainer must maintain his position as control­ler and apply the three Cs:

1. Communicate—the voice command tells him what he should do

2. Coordinate—apply the aids correctly to reinforce this command

3. Cohere—if you reward him appropriately, he will want to respond to you

As the horse becomes more confident and able, he will learn to discriminate more subtle signals than a voice com­mand. He will prepare to jump when we want him to even before we think we have asked him.

27.5.9.2.1 Behavior Chaining

Some tasks are more complicated and can be learned more efficiently by using a technique known as behavior chaining. It is most often used to teach complex behavior sequences, for example, in performing animals in the circus or in films. In these circumstances, we do not want to give a series of commands but want the animal to do a number of actions one after the other.

27.5.9.2.2 Punishment and Its Problems

Punishment is an integral part of the learning process but there are many problems with its use as a training aid. As a result, it tends to be misused and so causing a lot of concern. However, punishment is not a particularly efficient training tool, as it does not specifically tell the horse what it should be doing. It just signals that one specific activity in a given context should not be performed. Repeated use of punish­ment reduces its effectiveness and habituation occurs. Physi­cal punishment causes discomfort, inhibits learning, and if used excessively, emotional changes might occur which cause serious concerns such as aggression and learned help­lessness. Punishment makes a fearful horse worse and increases timidity.

27.5.10 Ethology of Wild Animals

It refers to studying the behavior of wild animals. Heini Hediger was the Swiss biologist and first scientist known as

the “Father of Zoo Biology,” who stressed the importance of ethology in captive wild animal management. The survival of a wild animal depends upon getting food, protection from other animals and adverse environment, and getting resis­tance to diseases, parasites, and predators. Wild animals’ behavior is species-specific and like other mammals, wild animals also exhibit specific pattern of behavior such as learning, feeding, sexual, social, communication, and agonis­tic behavior.

27.5.10.1 Communication Behavior in Wild Animals

Animals respond to external stimuli that come from signals given by other animals of their own species—from their parents or from their own offspring, for example, or from rivals or potential mates. Mammals use hearing, sight, and smell at a distance but for many of the more social ones that spend a good deal of time in physical contact, tactile commu­nication is also important. Chimpanzees touch and kiss each other’s hands in gestures of reconciliation after a fight. Animals that rely on vision for finding their way about and locating prey will also tend to communicate through visual signals. Sound has one big advantage in that it can go around corners and through some barriers, e.g., high-frequency (50-100 kHz) sounds are used by bats for echolocation. Animals can also help sound to travel further over longer distances by elevating themselves above ground. Crickets that sing from trees or shrubs can spread their signal over 14 times the area of those that sing from the ground and consequently attract more females. The territorial songs of birds are usually delivered from a raised song post which also increases their effective range, while grassland birds such as meadowlarks (Sturnella) and pipits (Anthus) sing while they fly.

In the animal kingdom, alarm calls are quite unique, e.g., monkeys give different alarm calls when they see different predators and chickens and ground squirrels also have differ­ent alarm calls for aerial and ground predators. Predator­specific alarm calls are particularly well-developed in vervets. When a vervet monkey sees a leopard (Panthera pardus) or other large cats, it gives a loud barking alarm call. When the other vervets hear this, they run up into trees so that the leopard cannot catch them. On the other hand, if the vervets catch sight of an eagle that prey on them they give a completely different alarm call—a sort of double-syllable cough. Since both these eagles are highly skilled at taking monkeys both from the ground and from the trees, a monkey on the ground hearing the eagle alarm call immediately looks up into the air and then runs into the thickest bush available. Yet another type of alarm call is given when the monkeys encounter snakes such as pythons or cobras. When a vervet hears a snake alarm call, it stands up on its hind legs and looks down into the grass.

Whales and elephants both use infra-sound to communi­cate over long distances. Elephants use low-frequency “rumbles” (10-35 Hz) that are very powerful and clearly effective over distances of 1-2 km to keep in touch with other members of their group. Some bats and owls also hunt in complete darkness not by emitting sounds of their own but by being extremely sensitive to sound made by their prey. The barn owl’s hearing is so good, for example, that it can locate a mouse in complete darkness simply by homing in on the sound of the mouse rustling through leaves or even just chewing.

In response to environmental stimuli such as visual, audi­tory, mechanical (touch), and chemicals (scent, taste, and pheromones) wild animals exhibit different type of behaviors. Among these behaviors feeding/eating, fly/run away, sexual, and maternal are considered as major behav­ioral responses. Aggressive behavior is more prevalent in wild animals. The major causes are competition of males for successful selection of females, fight over food resources, disturbance of social organization, incompatibility in the composition of groups, and distortion of social role.

Learning Outcomes

Significance of studying animal behavior: Behavior is closely related to the welfare of the animal. Hence, behavioral studies help the veterinarians and animal scientists to identify the cause of any abnormal behav­ior and address it. An understanding of the behavior of animals will guide the animal scientists to prevent and manage various behavioral disorders thus ensuring the welfare of the animal.

Endocrine moderation of behavior

Behavior is a complex response mediated by the nervous system and is modulated by the endocrine system. Hormones determine and influence the proba­bility that a specific sensory input leads to a specific behavioral response. Hormonal changes might modify some ongoing behavior by increasing or decreasing the frequency or duration of that behavior, or they might trigger the onset or end of a behavior or behavioral sequence.

Behavioral plasticity

The plasticity of behavior is an array of behavioral responses to varying environmental conditions. This determines the inherent potential of an animal to adapt to different environmental conditions. Broadly, three types of behavioral plasticity can be identified: differences in ontogenetic development, adjustments through learning, and the innate ability to respond to a variety of stimuli.

(continued)

Types of animal behavior

The behavior of animals has been broadly classified and studied extensively under categories such as learning, feeding, sexual, maternal, and social behaviors. Learning: Learning is a process whereby an individual acquires new responses and new capacities. Learning results in behavioral changes within an animal’s lifetime which can introduce a new dimension into behavioral evolution; Feeding behavior: is a circadian rhythm and an important sign of health which is influenced by a variety of internal and external factors; Sexual behavior: Sexual behavior is important in all species of animals. Sexual behavior includes proceptive and receptive behavior by the female and courting and mate guarding by the male, as well as actual copulation. Maternal behavior: The combination of the proper hormonal milieu and the stimulus for maternal behavior, the neonate, plus prior experience of being a mother can elicit maternal behav­ior. The stimulation of maternal behavior appears to be under both hormonal and neural control.

Wild animal behavior

It refers the studying the behavior of wild animals. Wild animals’ behavior are species-specific and like other mammals, wild animals also exhibit specific pat­tern of behavior such as learning, feeding, sexual, social, communication, and agonistic behavior.

Exercises

Objective Questions

Q1. The phenomenon wherein a young animal develops attraction to an object or an animal is called as

Q2. The type of communication which involves touch in behaviors such as social bonding, infant care, grooming, courtship, and mating is____________________.

Q3. The organization of a group of animals in such a way that some members of the group have greater access to resources like food or mates than others is called as

Q4. The area in the brain that accodes the level of priority to a given stimulus is.

Q5. The decrease in response to repeated or continuous stimulation is called as.

Q6. Pavlov conditioned dogs to salivate upon hearing a ringing bell by repeatedly presenting them with food while ringing a bell. This is an example of ______.

Q7. Coprophagy is seen in_________.

Q8. The fixed behavioral responses that animals exhibit even without learning them are known as

Q9. Stealing of young ones of other animals of their species by prepartum animals is seen in.

Q10. Vomeronasal organs are involved in detection of

Q11. Brooding behavior of the birds is influenced by

Q12. Group feeding animals tend to increase _________.

Q13. Punishing the animals during training leads to

Q14. Enurination is a unique sexual behavior seen in

Q15. Hormone that depresses feeding is _________.

Q16. Ensuing of abortion of the fetus when a strange male

mouse comes in contact with thepregnant female is known as.

Q17. One of the major hormones associated with maternal behavior is.

Q18. Peck order is a social ranking established in

Q19. One of the plastic changes induced by behavior in the brain components is.

Q20. In prey species such as cattle the most important mode of communication in disseminating information about general safety or danger is_____________.

Subjective Questions

Q1. Discuss the role of nervous system in development and exhibition of behavior.

Q2. Write briefly on the role of pheromones in animals.

Q3. Explain the control of feeding intake and the differences in feeding behaviors in animals.

Q4. Discuss the role of hormones on influencing the behavior.

Q5. Write a note on appeasing hormones.

Q6. Describe in detail the significance of social behavior.

Q7. Discuss in detail about the agnostic behavior observed in domesticated animals.

Q8. Write briefly on the various modes on social commu­nication in animals.

Q9. Write note on the types of learning.

Q10. Discuss in brief on the milking behavior and its factors controlling in cow.

Q11. Discuss in detail about the maternal behavior in animals.

Q12. Describe the various behavioral disorders related to sexual behavior.

Q13. Write briefly on the significance of wild animal ethology.

Q14. Discuss in detail the importance of ethology and role in animal welfare.

Q15. Describe the importance of training in domesticated animals and the commonly employed techniques.

Answer for the Objective Questions

A1. Imprinting

A2. Tactile

A3. Hierarchy

A4. Amygdala

A5. Habituation

A6. Operant conditioning

A7. Rabbits

A8. Instincts

A9. Ewes

A10. Pheromones

A11. Prolactin

A12. Feed intake

A13. Habitutation

A14. Goats

A15. Cholecyctokinin

A16. Bruce effect

A17. Oxytocin

A18. Chicken

A19. Synaptic organization and formation

A20. Vocal

Keywords for the Answers to the Subjective Questions

A1. Cerebral cortex, limbic system, hypothalamus, neural circuits

A2. Volatile chemicals secreted/excreted, vomeronasal organ, breeding activity, reducing aggression and fear, and encouraging feeding

A3. Short-term, long-term and emergency control, hormones, grazing, and ruminating behavior in cattle

A4. Influence during embryonic development, modify/ increase/decrease an ongoing behavior, influence social behavior

A5. Dams, maternal care, reduce aggressiveness, clinically used for behavioral problems

A6. Increased learning, and species-specific, survivability

A7. Threaten, attack to access limited resources, feed, terri­tory, and mating partner

A8. Vocal, olfactory, visual, and tactile communication

A9. Habituation, reinforcement, conditioning operant conditioning

A10. Neuroendocrine reflex, classical conditioning, oxyto­cin, stress, feeding concentrates

A11. Influence by hormone, hereditary, neural and experien­tial factors, nursing, grooming

A12. Lack of sexual interest towards receptive mares, self­mutilation, lack of ejaculation, masturbation and ten­dency to injure handlers, lack of socialization

A13. Species-specific, learning to communicate, alarm calls, searching for feed and water resources, social learning

A14. Understand the animal’s need, helps in better manage­ment, treatment, transport, and training

A15. Training for the basic daily life skills, used for live­stock handling, rescue dogs, assist visually impaired, therapy dogs, stress reduction purpose. Reinforcement, operant conditioning, reward

Further Reading

Atkinson T (2018) Practical feline behaviour understanding cat behaviour and improving welfare. CABI Publishing, Oxfordshire

Aubrey M, Dawkins M (2012) An introduction to animal behavior, 6th edn. Cambridge University Press, Cambridge

Barnard C (2004) Animal behaviour: mechanism, development, func­tion and evolution. Prentice-Hall, Harlow

Beaver BVG (2009) Canine behavior: insights and answers. Elsevier Health Sciences, St. Louis, MO

Broom DM, Fraser AF (2007) Domestic animal behaviour and welfare, 4th edn. CABI Publishing, Wallingford

Case LP (2010) Canine and Feline behavior and training: a complete guide to understanding our two best friends. Delmar, Cengage Learning, Clifton Park, NY

Dugatkin LA (2014) Principles of animal behavior, 3rd edn. W. W. Norton and Company, New York

Fraser AF (2010) The behaviour and welfare of the horse, 2nd edn. CABI Publishing, Wallingford

Houpt KA (2011) Domestic animal behavior for veterinarians and animal scientists. Wiley-Blackwell, Ames, IA

Jensen P (2007) Behavioural biology of dogs. CABI Publishing, Wallingford

Lethbridge E (2009) Knowing your horse: a guide to equine learning, training and behaviour. Wiley-Blackwell, Chichester

Phillips C (2002) Cattle behaviour and welfare, 2nd edn. Blackwell Science, Oxford

Price EO (2002) Animal domestication and behavior. CABI Publishing, Wallingford

Shaw JK, Martin M (2014) Canine and Feline behavior for veterinary technicians and nurses, 1st edn. Wiley-Blackwell, Chichester