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Cover
Title Page
Acknowledgments
About the Author
About the Companion Website
1 Communication
1. Introduction
2. Perception
3. Horses
4. Dogs
5. Cats
6. Pigs
7. Cattle
8. Sheep
9. Goats
2 Aggression and Social Structure
1. Sociability
2. Introduction
3. Categories of Aggression
4. The Biological Basis of Aggression
5. Cattle
6. Sheep
7. Goats
8. Horses
9. Pigs
10. Dogs
11. Cats
3 Biological Rhythms and Sleep and Stereotypic Behavior
1. Introduction
2. High‐Frequency Rhythms
3. Ultradian Rhythms
4. Circadian Rhythms
5. Other Rhythms
6. Annual Rhythms
7. Sleep Functions and Types
8. Patterns of Sleep and Activity in Domestic Animals
4 Sexual Behavior
1. Introduction
2. Cattle
3. Sheep
4. Goats
5. Horses
6. Pigs
7. Dogs
8. Cats
5 Maternal Behavior
1. Introduction: General Principles of Maternal Behavior
2. Pigs
3. Sheep
4. Goats
5. Cattle
6. Horses
7. Cats
8. Dogs
6 Development of Behavior
1. Introduction
2. Dogs
3. Cats
4. Horse
5. Pigs
6. Ruminants
7 Learning
1. Introduction
2. Comparative Cognition
3. Pigs
4. Dogs
5. Cattle
6. Sheep
7. Goats
8. Horses
9. Cats
8 Ingestive Behavior: Food and Water Intake
1. Introduction
2. Control of Food Intake in Pigs
3. Control of Food Intake in Dogs
4. Control of Food Intake in Cats
5. Control of Food Intake in Horses
6. Control of Food Intake in Cattle
7. Control of Food Intake in Sheep
8. Control of Food Intake in Goats
9. Water Intake
10. Specific Hungers and Salt Appetite
9 Behavioral Genetics
1. Cats
2. Dogs
3. Horses
4. Cattle
5. Sheep
6. Goats
7. Pigs
8. Laterality or Handedness in Animals
References
Index
End User License Agreement

List of Tables

Chapter 01
1. Table 1.1 The vocalizations of cats.
Chapter 03
1. Table 3.1 Percentage of time grazing by various populations of free‐ranging feral horses.
2. Table 3.2 Time budgets of stabled horses and ponies.
3. Table 3.3 Time budget of tethered horses.
4. Table 3.4 Time budget of Przewalski horses.
5. Table 3.5 Activity patterns of cattle in different environments.
6. Table 3.6 Mean values of comparative data of sleep–wakefulness states and attitudes in four species of farm animals (three subjects of each species).
7. Table 3.7 Activity patterns of sheep.
Chapter 07
1. Table 7.1 Results of imprint training.
2. Table 7.2 The types of learning that have been demonstrated in domestic animals.
Chapter 09
1. Table 9.1 Summary of canine fear and aggression GWAS results. Fear and aggression trait peaks are given for separate GWAS studies using Vaysse (marked “V”; Illumina HD) and Boyko (“B”; Affymetrix v.2) genotype datasets. Loci shared with both are black, and others are gray. Coordinates use CanFam2 assembly.

List of Illustrations

Chapter 01
1. Figure 1.1 Field of vision of the horse.
2. Figure 1.2 Greeting. Nostril‐to‐nostril investigation, in this case by a horse and pony.
3. Figure 1.3 The aggressive and submissive postures of horses. The ears of the aggressive horse on the right are back, while those of the submissive horse on the left are pointed sideways.
4. Figure 1.4 The mating expression of the mare.
5. Figure 1.5 The food‐anticipating expression of the horse.
6. Figure 1.6 The horse grimace scale.
7. Figure 1.7 Driving posture of the horse. The stallion, left, drives a mare. This behavior is also called snaking, herding, or driving (rounding).
8. Figure 1.8 The flehmen response, or lip curl. The location of the vomeronasal organ is indicated by the arrow.
9. Figure 1.9 Body postures of the dog. (A, B) Neutral to alert attentive positions; (C) play bow; (D, E) active and passive submissive greetings – note tail wag and shift in ear position and in distribution of weight on fore and hind limbs; (F–H) gradual shift from aggressive display to ambivalent fear‐defensive aggressive posture; (I) passive submission; and (J) rolling over and presentation of inguinal‐genital region.⁷²⁴
10. Figure 1.10 Elimination postures of the dog.²¹⁶²
11. Figure 1.11 Body postures of the cat. Aggressiveness is increasing from A₀ to A₃, fearfulness from B₀ to B₃. A₃ B₀ is the most aggressive cat, A₀ B₃ the most fearful, and A₃ B₃ the defensively aggressive cat¹³⁶⁰ from Overall¹⁷⁴⁵ with permission Elsevier.Facial expressions of the cat.¹³⁶¹ A₂ B₀ is offensively aggressive; A₀ B₂ is defensively aggressive. from Overall¹⁷⁴⁵ with permission Elsevier.
12. Figure 1.12 The gape expression of a cat. (A) The cat touches the investigated object with its nose and may lick its nose; then (B) it opens its mouth while gazing in a preoccupied fashion.
Chapter 02
1. Figure 2.1 Patterns of agonistic interactions in cattle. (A) Cows meeting after an active approach. The one on the left is threatening, whereas the one on the right has assumed a submissive posture. Note the head and leg positions of each cow. (B) Physical combat: a fight. The cows bunt or push against each other head to head, each striving for a flank position. (C) The clinch. One contestant of an evenly matched pair slips alongside the other; the head of the former is pushed between the legs and udder of the latter. In unusually prolonged contests, the cows rest briefly in the clinch between bouts of bunting. (D) Flank attack. The animal that gains a flank position is at a decided advantage over the other. The flanked animal either submits and flees or strives to regain the head‐to‐head position. (E) The butt. A dominant animal directs an attack against the neck, shoulders, flank, or rump of a subordinate, which in turn submits and avoids the aggressor. (F) Play fighting. The calf butts the mother.
2. Figure 2.2 An aggressive expression in a horse. A threat to bite.
3. Figure 2.3 Threat to kick. The horse on the right is threatening to kick the horse on the left. Note the lashing tail of the threatening horse and the tucked tail of the threatened horse.
4. Figure 2.4 Balking behavior by stallion on the right.
5. Figure 2.5 Agonistic behavior in boars. (A) Pawing the ground during initial encounter. (B) Strutting. (C) Shoulder‐to‐shoulder contact and slashing. (D) Perpendicular biting attack. (E) Submission of pig on the right. (F) Pursuit of the loser.
6. Figure 2.6 Grooming behavior of the pig. (A) The area scratched by the hind legs. (B) The area rubbed on vertical objects. (C) The area licked and nosed by other pigs.
7. Figure 2.7 Ranges used concurrently by three adult female (AP, BE, and SB) and one adult male (DL) cats in 1979. AP and BE shared three of four barns at the home farm (a) and most of the yards and pastures immediately surrounding the barns. SB was the only female at her farm (b). BE used the southern section of her range (c) through most of 1978 and 1979, and started using the northern area (d) in the late summer of 1979 in a series of foraging excursions with that year’s litter. One female kitten remained and eventually reproduced in this section of the range, whereas BE and a male kitten disappeared from this area in late 1979. The two section of BE’s range were connected only by the road between SB’s and AP’s ranges; BE hunted frequently along the road shoulders, and SB and AP foraged only in the adjoining pastures. AP and BE shared the area around an apartment complex (e), but never were noted to contact one another here. DL’s range included large areas of each female’s range, and on most evenings DL visited each barn complex at least once. The enclosed line at the lower end of the map represents 500 m. Immediately above it, the north–south axis is indicated. Dark quadrangles represent homes, apartments, and stores; open quadrangles, barns and other outbuildings. Narrow dotted lines indicate fence lines.
8. Figure 2.8 Grooming postures of the cat. (Top) Nongrooming. (Center) Flank grooming. (Clockwise) Grooming of hindquarters; scratching ears; the first step of face grooming, licking the front leg. Solid arrows indicate grooming sequences of normal cats. Dotted and dashed arrows indicate sequences in cats with tectal lesions.
Chapter 03
1. Figure 3.1 Nyctohemeral cycle of melatonin concentration in calf plasma (upper) and cerebrospinal fluid (lower) at selected times of the day. The lights were on from 0600 to 2000 hours. Data points are means ±1 standard error of samples from number (N) of calves. Letters indicate that when two means do not have any letter in common, they are significantly different.
2. Figure 3.2 Circadian rhythms of corticosteroid secretion in pigs. The graph is based on the mean results from six pigs. The peak of corticosteroid output occurs in late morning.
3. Figure 3.3 The stages of vigilance and sleep in the cat. Polygraphic record at the speed of 30 mm per second showing the stages identified. LO‐RO: electroencephalographic record from the left and right occipital area; LF‐RF: electroencephalographic record from the left and right frontal area.
4. Figure 3.4 Time budgets of domestic herbivores. Grazing, standing, and lying behavior of cattle, sheep, and horses living on the same pasture during two seasons.
5. Figure 3.5 Example of the activity pattern of a dog recorded with an accelerometer. Note the diminished activity between 2300 and 0630 hours and the peak of activity in the evening, presumably when the owner returned home from work.
6. Figure 3.6 (A) Physiological parameters of sleep in various species. The different electrocorticogram (EcoG) patterns for each species are shown at a speed of 15 mm per second: theta rhythm (horse), delta rhythm (ruminating cow), spincles (sheep), and alpha rhythm (pig). (B) Mean comparative data of sleep and wakefulness states and of postures during nighttime. The inner circle shows the relative duration of the EcoG pattern (rapid eye movement [REM] in dark green), and the outer circle shows the relative duration of the postures.
7. Figure 3.7 The postures of the horse when lying down and getting up. (A,B) Lying down; (C,D) getting up; (E) sternal recumbency: the horse not lying symmetrically but with the lateral surface of one foreleg on the ground; (F) lateral recumbency: the upper foreleg anterior to the lower one. The horse exhibits REM sleep in (F) or in (E) with the muzzle touching the ground.
8. Figure 3.8 Mean frequency of cribbing before and after delivery of roughage meal (upper graph) and concentrated meal (lower graph). Dotted line indicates the mean level. *P < .05; **P < .011287.
9. Figure 3.9 Comparison of monthly variation in patterns of activity budgets of male and female adult goats on Rum in (A) 1981 and (B) 2000. Only the proportions of daytime spent feeding and in other activities are shown.
Chapter 04
1. Figure 4.1 Factors that affect sexual behavior.
2. Figure 4.2 Relationship of estrogen, progesterone, and LH levels to behavior in the bitch. Times of onset of behavior and physical signs represent means of a range of times.³⁷⁷
3. Figure 4.3 Effects of testosterone (T), estradiol (E₂), estrone (E₁), and dihydrotestosterone (DHT) on agonistic behavior, vulvar interest, the flehmen response, and mounting.⁷¹³
4. Figure 4.4 Average composition of daily activities of cattle during estrous and non‐estrous stages of the reproductive cycle.⁹¹¹
5. Figure 4.5 Mounting of cows at different stages of estrus by cows and bulls.
6. Figure 4.6 The mean (± SEM) response of bulls to estrous and non‐estrous urine, mucus, and water.
7. Figure 4.7 Sexual patterns in cattle, sheep, and goats.
8. Figure 4.8 The occurrence of masturbation by time of day. The masturbations observed are plotted by the hour of the day in which they occurred as the percentage of each bull’s masturbations. Some bulls were observed longer than others, so it was necessary to use percentages rather than absolute numbers of masturbations.¹⁰⁴⁵
9. Figure 4.9 Response of the ram to urine of an estrous ewe. (A) Urination by ewe; (B–D) ram nosing urine on the ground; (E) the Flehmen response.
10. Figure 4.10 Nudging sequence in sheep courtship behavior.
11. Figure 4.11 (A) Schematic diagram of a coronal section through the sheep at the level of the optic chiasm (OC) and anterior commissure (AC). The position of the ovine sexually dimorphic nucleus (oSDN) is shown bilateral to the third ventricle (V) in the central part of the medial preoptic nucleus (MPOA). The ventral paraventricular nucleus (vPV) is shown at the base of the ventricle. (B) Micrograph of oSDN from the left hypothalamus of a female‐oriented ram. The third ventricle is at the right of the figure. (C) Section of a male‐oriented ram comparable to that in (B). The photomicrograph is taken near the midpoint of the anterior–posterior extent of the oSDN. The scale bar is 1 mm.
12. Figure 4.12 (A) The posture of the estrous mare. The clitoris is everted, and the tail deviated. (B) The prance, or piaffe, of the courting stallion.
13. Figure 4.13 The courtship sequence of pigs.
14. Figure 4.14 Copulatory lock in dogs.
15. Figure 4.15 Percentage of dogs experiencing rapid decline, gradual decline, or no change in four behaviors after castration.
16. Figure 4.16 Sexual behavior of the female cat. (A,B) The typical posture adopted by the queen in full estrus. Note leg flexion, lordosis, and deflection of the tail. (C) Tom holding queen with neck grip during intromission. (D) Postcoital rolling by the queen.
Chapter 05
1. Figure 5.1 Factors that influence the expression of maternal behavior. The horse is used as an example, but there are similar influences on maternal behavior in all domestic animals.
2. Figure 5.2 Ewe licking the head of her newborn lamb.
3. Figure 5.3 Initial orientation of the lamb to the ewe’s underline, but in the axilla rather than the udder.
4. Figure 5.4 The bond of the lamb to its mother is stimulated by colostrum ingestion, which leads to release of cholecystokinin and stimulation of vagal afferents.
5. Figure 5.5 Changes with age in frequency of nursing by pony foals.
6. Figure 5.6 Identification of the foal. The mare sniffs the anal area of the foal. The hind leg on the side opposite the foal is flexed to facilitate nursing.
7. Figure 5.7 The effect of masking vision, olfaction, or both on the time that a foal takes to locate its dam.
8. Figure 5.8 Approach of the queen to her kittens and kittens to the queen during the first eight weeks of life.
9. Figure 5.9 The decrease in licking of the puppies by the bitch as the puppies mature.
Chapter 06
1. Figure 6.1 Behavioral development of the dog. Superscript numbers refer to the references: (1) ^723,730, (2) ¹⁹²⁹, (3) ²³⁶, (4) ²⁶³, (5) ²⁰⁶⁶, and (6) ^726,728.
2. Figure 6.2 The average number of vocalizations by puppies at different ages and in two environments. Note that vocalization of a puppy alone in the home pen is consistently lower than that of one in a strange pen. Puppies whose tests were begun when they were older than three weeks showed a slightly higher rate of vocalization, but the curves are parallel.¹⁶⁶⁷
3. Figure 6.3 Behavioral development of the cat. Superscript numbers refer to the references: (1) ¹²³⁴, (2) ¹⁹⁷⁴, (3) ⁷²⁵, (4) ⁷²², (5) ¹³⁶¹, (6) ²⁴²⁶, (7) ²⁰⁷⁸, (8) ⁴⁴¹, (9) ²³⁵³, and (10) ²²⁹⁸.
4. Figure 6.4 The change in playing behavior of kittens with age. Social play reaches a peak at 12 weeks and then declines. Frequency refers to the percentage of the daily 90‐minute observation period in which play was observed.²⁰⁰³
5. Figure 6.5 Behavioral development of the horse. References: ^{239,480,483,484,593,2386}.
6. Figure 6.6 The mean percentage of time the foals spent feeding when their mother was feeding, active, or resting upright.
7. Figure 6.7 Relative frequency of various types of play. The mean percent of total play bouts by fourteen foals in which the type of play was running and bucking alone, running and bucking as a group, interactive, manipulation of an object, or play at an adult. Standard error of the mean is shown as a vertical bar. (With permission Dr. Sharon Crowell‐Davis).
8. Figure 6.8 Relative frequency of various types of play. The mean percentage of total play bouts by 14 foals in which the type of play was running and bucking alone, running and bucking as a group, interactive, manipulation of an object, or play with an adult. Standard error of the mean is shown as a vertical bar.⁴¹³
9. Figure 6.9 Submissive snapping of the immature horse. Foal approaches its dam while snapping.²⁴⁸⁴ Rate of flehmen by foals of various ages. Colts exhibit flehmen more than fillies only during the first few weeks. Asterisks indicate significant difference.⁴⁸¹
Chapter 07
1. Figure 7.1 Classical conditioning. Pairing the sound of the metronome with shock to the foreleg conditions a goat to lift its leg when a metronome ticks. Also shown is the late Professor Liddell, who compared the rate at which various species of farm animals learned a classical conditioning.
2. Figure 7.2 Operant conditioning. The dog pushes a pedal (operates on the environment) to obtain a dish of food.
3. Figure 7.3 The relative brain size of various domestic animals. Right row, from top to bottom: horse, cow, and pig; left row: dog, sheep, and cat.
4. Figure 7.4 Canine cognitive bias test. (A) Owner holds dog in preparation for training session. (B) Dog learning that the bowl on the right always contains food. (C) Dog approaching the bowl on the left by a circuitous route. He has learned that there will be no food there. (D) The cognitive bias test. The speed of the dog approaching a bowl midway between the always‐rewarded side and the never‐rewarded side is a measurement of his optimism.
5. Figure 7.5 Equine cognitive bias test. (A) Horse learning that the bowl on the right will never contain feed. (B) Horse approaching the feeder on the left that always contains food. He has learned that there will always be food there. (C) Handler holds horse in preparation for cognitive bias test. (D) The cognitive bias test. The speed of the horse from the time she is released and the rope is dropped until she reaches the feeder midway between the always‐rewarded side and the never‐rewarded side is a measurement of her optimism.
6. Figure 7.6 Learning in neonatal dogs. The puppy learns to avoid the textured side, for which puppies have an innate preference, if that side is associated with a blast of air.
7. Figure 7.7 An example of a learning deficit associated with organic disease of the central nervous system, showing the brain of a Lhasa apso that suffered from lissencephaly and could learn neither bowel nor bladder control.
8. Figure 7.8 Taste aversion learning in cattle. On day 1 (trial 1), the experimental cows (open symbols) were offered alfalfa pellets. After they had eaten the food, apomorphine was injected. The control cows (closed symbols) were injected with 0.9% NaCl. Four days later (trial 2), when all cows were offered alfalfa pellets, the cows that had been treated with apomorphine ate almost nothing, whereas the controls ate more than they had initially. The experimental cows learned to avoid a feed associated with illness.
9. Figure 7.9 A horse demonstrating that he understood the concept of larger by choosing the largest symbol seven years after he was originally taught the discrimination.
Chapter 08
1. Figure 8.1 Integration of physiological factors that stimulate and depress food intake in the brain.
2. Figure 8.2 Integration of pathological factors that depress food intake.
3. Figure 8.3 Daily food intake of pigs fed ad libitum. Pig on left was subjected on alternating days to temperatures of 25 °C (white) and 10 °C (green). Pig on right was subjected on alternating days to temperatures of 25 °C (white) and 35 °C (green).
4. Figure 8.4 Effect of various gastric loads on three‐hour intake of suckling pigs. Note that milk (Similac) and isotonic glucose, but not isotonic saline, depressed intake.¹⁰⁵⁰
5. Figure 8.5 Effect of hypertonic injections administered into the duodenum (left), the jugular vein (center), or the portal vein (right) on subsequent food intake of pigs. Only intraduodenal injections depressed intake relative to saline‐injected controls.
6. Figure 8.6 Intake of a pig following injection of saline (white) or insulin (green).
7. Figure 8.7 Effect of novelty on food preferences. Dogs normally prefer diet P to diet R/CC, but will show preference for either R/CC or P if it is a novel diet. Preference for P is sustained (lower graph), but that for R/CC is reversed in a week (upper graph).
8. Figure 8.8 Effect of ovariohysterectomy on food intake and body weight of Beagle bitches. Solid lines = intact. Dashes = spayed bitches.
9. Figure 8.9 Food intake and body weight of two Beagles. Ad libitum feeding began at arrow. Lower graph is of a dog whose intake and body weight remained normal. Upper graph is of a dog that overate and became obese.
10. Figure 8.10 Intake of two horses following intravenous saline (dark) or diazepam (light).
11. Figure 8.11 Reciprocal relationship between dietary characteristics and dry matter intake. Line a to b represents intake limited by the fill effect of the diet. Line b to c represents intake limited by the energy demand of the animal. Dashed lines represent unattained intake predicted by extrapolating the theoretical equations.
12. Figure 8.12 Controls of feeding in a ruminant.
13. Figure 8.13 Changes in daily NaCl intake by ponies after furosemide or CON treatment (mean standard error of the mean).
Chapter 09
1. Figure 9.1 Breed differences in feline affection to human family members.
2. Figure 9.2 Breed differences in feline aggression to other cats.
3. Figure 9.3 Breed differences in various types of aggression in dogs.
4. Figure 9.4 Behavioral independence in Tennessee Walker mares varies with agouti signaling protein (ASIP) loci. Black mares (aa) (on the right) are more independent (self‐reliant) than bay mares (A−) (on the left).
5. Figure 9.5 Percentage distribution of complete copulations, mounts without penis erections, and mounts without thrusts in pairs of twin bulls.
6. Figure 9.6 Avoidance learning in pigs. (A) To avoid shock, the pig must jump a barrier when a buzzer sounds. (B) The ability to learn to avoid shock decreases with age in Duroc and Hampshire pigs.

This edition first published 2018
© 2018 John Wiley & Sons, Inc.

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Iowa State University Press (4e, 1982); John Wiley & Sons, Inc. (5e, 2011)

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Library of Congress Cataloging‐in‐Publication Data

Names: Houpt, Katherine A., author.
Title: Domestic animal behavior for veterinarians and animal scientists / Katherine A. Houpt.
Description: 6th edition. | Hoboken, NJ : John Wiley & Sons, Inc., 2018. | Includes bibliographical references and index. |
Identifiers: LCCN 2017051648 (print) | LCCN 2017053200 (ebook) | ISBN 9781119232797 (pdf) | ISBN 9781119232803 (epub) | ISBN 9781119232766 (hardback)
Subjects: LCSH: Domestic animals–Behavior. | Animal behavior. | MESH: Animals, Domestic | Behavior, Animal
Classification: LCC SF756.7 (ebook) | LCC SF756.7 .H68 2018 (print) | NLM SF 756.7 | DDC 636.089–dc23
LC record available at https://lccn.loc.gov/2017051648

Cover Design: Wiley
Cover Image: Courtesy of Charles E. Houpt