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4 Management of Breeding Colonies REPRODUCTION To maintain the breeding efficiency of a colony or to breed an impor- tant individual dog successfully, staff must understand the unique reproduc- tive characteristics of dogs. The biology of canine reproduction has been extensively reviewed (Burke, 1 986; Christiansen, 1 984; Concannon, 1 99 1; Concannon and Lein, 1989; Concannon et al., 1989~. Information on herita- bility of physical and other characteristics of dogs, Mendelian genetics of breeding, the incidence and characteristics of diseases that have a genetic basis, and methods for demonstrating heritability is also available (Patterson, 1975; Patterson et al., 1989; Shultz, 1970; Willis, 1989~. Reproductive Cycle of the Bitch Most bitches can become pregnant once or twice a year. Each ovarian cycle consists of the following phases: . A follicular phase, or proestrus, during which there is progressive vulval swelling and a serosanguineous (bloody) vaginal discharge. During this period, which can last from 3 days to 3 weeks, the bitch's blood has high concentrations of estrogen. The male will show interest, but he either does not or is not allowed to mount. . A periovulatory period, or estrus, during which estrogen declines 35

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36 DOGS: LABORATORY ANIMAL MANAGEMENT and progesterone increases as the ovarian corpora lutea form This period is also the early luteal phase of the cycle. During estrus, which can last from 3 days to 3 weeks, the bitch assumes a characteristic posture in the presence of a male in which the rump is raised and there is a curvature of the back (lordosis) and the tail is held to one side (flagging). The male is allowed to mount, and copulation occurs. A midluteal and late luteal phase or metestrus (either pregnant or nonpregnant metestrus', which lasts about 2 months and during which se- rum progesterone remains elevated above 1 ng/ml. A period of weak ovarian activity, or anestrus, lasting 2-10 months, in which progesterone concentration is low, and there is no evidence of estrogen stimulation of the uterus or vulva. In constant photoperiods of 12 hours of light and 12 hours of darkness or 14 hours of light and 10 hours of darkness, estrous periods should occur with equal incidence throughout the year. Possible effects of constant light With natural circannual changes in photoperiod, bitches come into estrus more frequently in winter and spring months than in summer and autumn months. In most breeds, the interval between es- trous periods averages 7-8 months. After the age of 8 years, however, the interval between cycles begins to lengthen, reaching 12 months or longer by the age of 12 years (Andersen and Simpson, 1973~. Successful breeding requires that observation of reproductive condi- tions be given high priority, and staff must be able to recognize the start of proestrus. A swollen vulva might not be obvious on a dark or long-haired dog, and bitches often lick away the bloody discharge; therefore, the vulva of each breeding bitch must be examined closely two or three times a week, beginning 4 months after estrus. Vaginal cytology can be useful for estimating the best time for breeding (Concannon and DiGregorio, 1986; Hoist, 1986; Olson et al., 1984) and predicting the time of whelping, which will be 55-60 days after a change in the smear indicates late estrus. Vaginal smears in anestrus are nondescript, with a few leukocytes and small epithelial cells. In early proestrus, smears include a high proportion of rounded epithelial cells, erythrocytes, and sometimes a few leukocytes. During midproestrus, there is an increasing percentage of cornified (flakelike) epithelial cells but no leukocytes. All or nearly all epithelial cells in the smear are cornified from 2-8 days before ovulation until 4-9 days after ovulation, when these cells predictably and abruptly decline. In early metestrus, cornified cells are replaced by rounded, smaller superficial cells, and there is usually an influx of leukocytes. The metestrus smear slowly regresses to the nondescript anestrus smear. Smears should be taken from the anterior vagina. They should be obtained and prepared carefully with saline-moistened swabs and should not be contaminated with have not been studied.

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MANAGEMENT OF BREEDING COLONIES 37 vulval material. In the case of bitches that have had reproductive problems, when a successful breeding is important, or both, more accurate predictions can be made by monitoring the progesterone concentration in the serum or plasma with an enzyme-linked immunosorbent assay (ELISA) kit (Bouchard et al., 1991a; Hegstad and Johnston, 1992; Johnston and Romagnoli, 19914. In this test, ovulation occurs a mean of 1-2 days after the initial rise in progesterone, peak fertility a mean of 0-4 days after the initial rise, loss of fertility 6-11 days after, implantation 18-20 days after, and parturition 63- 65 days after (Concannon, 1991~. Mating Theoretically, it is sufficient to maintain one male for every 10-20 females; however, in practice this ratio might not be adequate, for several reasons. First, a bitch in proestrus produces pheromones that will start proestrus in other bitches in the colony, making it likely that several bitches will be in estrus simultaneously. Because mating an individual male more often than once each day can reduce its sperm output after 1 week (Amann, 1986), a greater ratio of males to females might be required to maintain breeding efficiency. Second, except under special circumstances, such as reproducing a disease model, breeding programs should conscientiously avoid inbreeding, and it has been estimated that a ratio greater than two males for each 10 females is needed to prevent an increase in the coefficient of in- breeding (Shultz, 1970~. Natural Mating Mating can be done naturally or by artificial insemination with fresh or frozen and thawed semen. Provided that the male is healthy, it is not necessary to take special precautions or to use medications to treat the genitalia because the vagina is not a sterile environment. However, it is important to ascertain that neither the dog nor the bitch has canine brucellosis, a disease that seriously affects reproduction and is a zoonosis (see "Control of Infectious Diseases" in Chapter 5~. The bitch is usually taken to the stud dog's pen or cage, because a dog will often ignore the bitch or spend an inordinate amount of time scent-marking if he is moved to new surround- ings. The bitch should be mated on 2 or 3 days over a 3- to 5-day period. Unless the staff is experienced in distinguishing early proestrus from estrus, the bitch should be presented to the male for 10-15 minutes every day or every other day from the time she is found to be in proestrus until she is mated. Breeding pairs should not be left unattended, because some bitches are highly selective in choosing mates and it is not uncommon for a bitch to attack a dog that is not of her choosing. In addition, the dog might need

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38 DOGS: LABORATORY ANIMAL MANAGEMENT assistance until he attains a copulatory lock. Mating should be recorded only on the basis of observations of copulatory locks that last several min- utes or more. If a bitch refuses a particular dog, even when signs of estrus (lordosis and flagging) are present, placing her with a different dog might solve the problem. If it is important to the breeding program that a bitch be bred to a dog that she is refusing, a caretaker should restrain her in a manner that will prevent her biting either the caretaker or the stud dog during breeding, or artificial insemination (AI) should be used. To ensure accuracy of parentage, the same stud must be used for every breeding within a single estrus to avoid multiple-sire litters. Bitches allow dogs to mate from several days before ovulation until several days after ovulation. Because the events of pregnancy are related to the time of ovulation not necessarily to the time of mating parturition can occur 56- 68 days after a single mating and up to 70 days after the first of multiple matings. Sperm can survive 6 days or more in the bitch, and ovulated eggs can remain fertile for 3-7 days. Parturition should occur 62, 63, or 64 days after ovulation in nearly every bitch (Concannon et al., 19831. Bitches that whelp 56-60 days after the first mating often have small litters, probably because they were bred at the end of the fertile period (P. Concannon, New York State College of Veterinary Medicine, Cornell University, Ithaca, N. Y., unpublished). Artificial Insemination AI can be helpful when males cannot be moved easily within or be- tween facilities, when breeding females with weak or selective estrus be- havior, when using males that cannot provide natural service, and for pre- serving valuable animal models. Semen collection, handling of semen, and insemination are described in detail elsewhere (Christiansen, 1984; Concannon and Battista, 1989~. Insemination with fresh semen. Semen can be collected in a clean paper cup or in a latex cone (artificial vagina) attached to a 15-ml conical polypropylene centrifuge tube. An advantage of the former method is that debris from the penis is less likely to become mixed in the ejaculate. Ejacu- late should be maintained at room or skin temperature and should be checked microscopically for sperm viability and malformations. Any variation from the expected chalky white color or 1- to 5-cc volume should be recorded. The full ejaculate should be deposited into the anterior vagina with a clean plastic pipes attached to a syringe with nonrubber (e.g., polypropylene) tubing. The hindquarters of the bitch should be raised for 10 minutes while the vagina is manipulated digitally by an attendant wearing a clean glove. The bitch should not be allowed to sit for 20 minutes, and pressure on her

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MANaGEMENT OF BREEDING COLONIES 39 abdomen should be avoided. AI should be performed every other day until two or three inseminations have been accomplished. The precise timing for performing AI can be predicted by checking for softening of the vulva, which often occurs around the time of ovulation; by demonstrating the appropriate vaginal cytologic characteristics of advanced estrus; or by mea- suring the initial rise in serum or plasma progesterone. Ideally, two insemi- nations should occur before vaginal smears show reduced cornification. Insemination with fresh chilled semen. Fresh semen can be diluted or extended in one of several laboratory buffers or commercial extenders and shipped refrigerated by overnight express for use in insemination in another location (Concannon and Battista, 19891. At 4C (39.2F), sperm motility remains nearly normal for 3-4 days if the semen is diluted in an appropriate diluent and for 1 day if undiluted (see Morton and Bruce, 1989~. Insemination with frozen semen. Frozen semen should be thawed and handled according to the instructions provided by the laboratory that pro- cessed it, because each freezing technique has stringent requirements for rate of thawing, dilution, and site of deposition. Although sperm live for several days in fresh semen, they normally die within a few hours after thawing; therefore, precise timing of insemination is important for success- ful impregnation. The best time to inseminate is usually shortly after oo- cyte maturation, which occurs 5-6 days after the initial rise in progesterone, around the time of a surge in leutinizing hormone. In most bitches, the inseminations should also take place 2-4 days before the decrease in vaginal cornification. Reported success rates for vaginal insemination range from O to 70 percent (Concannon and Battista, 1989~; success probably depends heavily on the freezing method and the number of viable sperm insemi- nated. Success rates of 50-90 percent have been reported for uterine in- semination, which is accomplished surgically or with special instrumenta- tion to deposit sperm through the cervix (Concannon and Battista, 1989~. Pregnancy and Parturition Pregnancy can be determined at 25 days after ovulation by ultrasonography, at 20-35 days after ovulation with palpation, and at 45 days after ovulation with radiography (Johnson, 1986; Yeager and Concannon, 1990~. There are no well-documented biochemical or immunologic canine pregnancy tests available. Concannon (1991) has reviewed changes in body weight during pregnancy and pregnancy-specific changes in hematocrit, serum chemistry, and metabolism. Whelping facilities should provide seclusion from excessive noise and other disturbances. The whelping box should be large enough to accommo

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40 DOGS: LABORATORY ANIMAL MANAGEMENT date the bitch and pups and have sides high enough to prevent neonates from wandering out of the box. The bottom of a large, fiberglass shipping crate works well for beagle-size dogs. The whelping box should be pro- vided about a week before expected parturition. Johnson (1986) has reviewed the management of the pregnant bitch. A nonpurulent green discharge, anorexia, and restlessness are normal just be- fore parturition. Birth of a litter can be either rapid or protracted over much of a day. Intervals between pups normally range from 20 minutes to 3 hours. Intervals greater than 3 hours can indicate a problem with fetal position or uterine function and warrant veterinary attention. Persistent, unproductive labor of more than 1 hour also requires veterinary attention (Johnston and Romagnoli, 1991; Jones and Joshua, 1988~. NEONATAL CARE Newborn pups, like all neonatal mammals, have poorly developed tem- perature-control mechanisms; therefore, it is necessary to keep the tempera- ture in the whelping box higher than room temperature. Temperatures of 29.4-32.2C (85-90F) have been suggested for the first 7 days of life, 26.7C (80F) for days 8-28, 21.1-23.9C (70-75F) for days 29-35, and 23.9C (70F) thereafter (Poffenbarger et al., 19901. That can be done by raising the temperature of the room and placing insulation between the whelping box and the cage or floor or by using heating devices, such as heat lamps or built-in heating elements. However, caution is necessary in using such heating devices; because pups younger than 7 days old have very slow withdrawal reflexes (Breazile, 1978), they can be overheated or severely burned by these devices. Circulating-water heating pads or commercial pig warmers are useful, because they maintain heat at a safe level. Whelping boxes should be examined two or more times a day for evi- dence of maternal neglect or cannibalism and for problems with the pups. A normal pup is plump and round, its head is mobile, and it exhibits a rooting reflex. Breathing is regular and unlabored, and the coat is shiny and free of debris. Abdominal enlargement after nursing is normal, but abdomi- nal enlargement accompanied by restlessness, weakness, and either exces sive vocalization or complete silence can indicate illness or aerophagia. Failure to gain weight is often the first sign of illness in a newborn animal (Greco and Watters, 19901. Andersen (1970) reported expected weight gains for beagle pups. Dead pups should be removed from the box. Andersen (1970) and Lawler (1989) have reviewed causes of neonatal deaths and have reported an average rate of death of about 20 percent. Necropsy examination is suggested for all pups that die or are euthanatized with severe illness. Such examinations are necessary to distinguish between congenital defects, which affect only the pups in which they occur; infectious diseases, whose spread

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MANAGEMENT OF BREEDING COLONIES 4 l might be prevented; and problems with the dam (e.g., insufficient milk) or the environment (e.g., room temperature too low), which can be corrected. REPRODUCTIVE PROBLEMS False Estrus and Anestrus Recurrent frequent false estrus (estrus without ovulation) has been re- ported (Shille et al., 1984~. In false estrus, estrus appears normal, and bitches will mate but fail to conceive. False estrus can be confirmed by demonstrating with a progesterone ELISA kit that the serum or plasma progesterone concentration has not risen above 1 ng/ml, as would be ex- pected for 50 days or more after ovulation if estrus were normal. Bitches that often have false estrus or have false estrus followed in a few weeks by normal estrus cause problems in maintaining breeding colonies. Except in special circumstances, such as reproducing a disease model, it is preferable to cull these animals. Culling based on small litter size, problems with whelping or maternal behavior, chronic infertility, or persistent anestrus is also appropriate. Methods for assessment and treatment for potential causes of infertility in females have been extensively reviewed (Feldman and Nelson, 1987; Johnston and Romagnoli, 1991; Shille, 1986~. Persistent anestrus can be distinguished from unobserved cycles only through extremely careful examinations for signs of proestrus or progesterone assays every 6 weeks. Estradiol assays are not particularly informative, and assays of canine go- nadotropin to diagnose primary gonadal failure are not readily available. Attempts to induce estrus in anestrus bitches have had variable success (Bouchard et al., l991b; Concannon, 1992, Concannon et al., 1989~. Delayed Parturition Whelping should not be considered overdue until 67 days after the last mating or possibly 70 or more days after the first of several matings. Ce- sarean section should not be contemplated earlier unless there are obvious signs of distress in the bitch. Johnson (1986) and Jones and Joshua (1988) have reviewed veterinary management of dystocia. Pseudopregnancy Bitches that are not bred or that are bred but fail to become pregnant frequently exhibit pseudopregnancy because of the progesterone secretion that always follows ovulation. Signs of pseudopregnancy include extensive mammary development, lactation, and maternal behavior. Pseudopregnancy is rare in beagles but more common in other breeds. It is self-limiting and usually does not require intervention (Feldman and Nelson, 1987~.

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42 DOGS: LABORATORY ANIMAL MANAGEMENT SPECIAL NUTRITIONAL REQUIREMENTS Bitches During pregnancy and lactation, bitches should be fed a diet approved by the Association of American Feed Control Officials for all life stages or a diet specially formulated for gestation and lactation (see "Selecting Opti- mal Rations" in Chapter 3~. When a quality diet is fed, supplementation with vitamins and minerals is neither necessary nor desirable. During the first two-thirds of pregnancy, the amount fed should be the same as that fed before pregnancy. During the last trimester, food intake should be gradually increased so that at parturition it is 150 percent of the daily maintenance requirement. Bitches should not be permitted to become obese during gestation, because this condition can increase the risk of dystocia and postparturient metabolic disorders (Johnston, 1986~. Bitches that are underfed during gestation tend to have a higher incidence of stillbirths than bitches that are fed appropriate amounts, and their pups often weigh less at birth (Holme, 1982~. Lactation represents the greatest nutrient challenge that bitches experi- ence during their lifetimes. For the first 3 weeks after parturition, nutrient requirements increase rapidly, leveling off at 200-250 percent of daily mainten- ance requirements, or even more, depending on the number of nursing pups (NRC, 1985~. The nutritional demands of lactation are met best through free access to both food and water. At the time of weaning, food is gener- ally withheld for 24 hours to decrease milk production. Food intake for the first day after weaning should be one-fourth of the amount required for maintenance and then gradually increased to the maintenance requirement by day 4. Ideally, lactating bitches should be within 15 percent of their prebreeding body weight at the time of weaning (AAFCO, 1993~. Pups Pups should be maintained exclusively on their dams' milk until they are 3 weeks old. They can then begin to eat small amounts of a moistened gestation-lactation diet or a growth diet. Most pups can be weaned com- pletely onto this type of diet by the age of 6-8 weeks. For the development of normal social behavior, it is desirable that they not be completely weaned before they are 6 weeks old. Pups that cannot be nursed by their dam or a foster dam before they are 5 weeks old should be fed one of the commer- cially available, complete milk replacers. Pups can be fed with bottles and nipples or stomach tubes. Bottles and nipples should be thoroughly cleaned after each use. If a stomach tube is used, its proper placement can be

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MANAGEMENT OF BREEDING COLONIES 43 ensured by inserting it to a distance equal to the premeasured distance from the mouth to the last rib. A small amount of sterile saline solution should be introduced through the tube before milk replacer is injected. After each meal, orphaned pups should be massaged in the anal-genital region with a warm, wet cotton ball to stimulate urination and defecation. Most orphans can be completely weaned onto solid food by 5 weeks of age. Young pups most readily eat canned or moistened dry food; older pups can be fed dry, semi-moist, or canned food. Pups can be fed on either a free-choice or meal-feeding program. If a meal-feeding program is used, they should be fed at least four times a day until they are 3 months old, three times a day until they reach two-thirds of their adult weight, and two times a day thereafter. After the age of 3 months, free-choice programs can lead to obesity in small breeds and faster than optimal growth in large breeds. Excessively rapid growth in breeds whose weight at maturity is more than 30 lb has been associated with an increase in the incidence of several metabolic bone diseases (Hedhammer, 1981; Hedhammer et al., 1974; Kealy et al., 1992~. Pups should be fed so that they grow at near optimal rates; growth-curve data are often available from pet-food manufacturers. When an appropriate growth ration is fed, no supplementation is necessary. If a product is not capable of supporting an optimal growth rate, it is gener- ally safer, less expensive, and more convenient to switch to a better-quality growth diet. As a general rule, pups gain approximately 1-2 g/day per pound of anticipated adult body weight (Lewis et al., 1987~. An inappropri- ate growth rate usually reflects a problem with the ration being fed or with the pups' access to it. VACCINATION AND DEWORMING Annual vaccinations and deworming of brood bitches should be sched- uled for anestrus or weaning periods, not when bitches are in proestrus or are pregnant. Pups that have nursed on colostrum during the first 12 hours after birth have received passive immunity to viruses against which the dam was im- munized. If pups cannot nurse on colostrum, 16 ml of pooled serum admin- istered subcutaneously has been shown to be a successful alternative (Bouchard et al., 1992~. Maternally acquired immunity declines over time, and the rate of decline, although variable, depends on the level of the dam's immunity at parturition and the amount of colostrum ingested by each pup. About 30-50 percent of pups will be susceptible to disease and capable of being effec- tively vaccinated by the age of 6-7 weeks. Most pups (more than 95 per- cent) can be effectively vaccinated by the age of 16 weeks. General prin- ciples of immunity in newborn animals and of immunoprophylaxis are reviewed elsewhere (Carmichael, 1983; Tizard, 1977a,b). Diseases to which pups are

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44 DOGS: LABORATORY ANIMAL MANAGEMENT susceptible and vaccination schedules are discussed in Chapter 5, Veteri- nary Care. Roundworms (Toxocara cants) and hookworms (Ancylostoma caninum and A. braziliense) are endoparasites that commonly infect young pups. Roundworms are typically transmitted from bitches to pups in utero, and pups begin to shed eggs in their feces 3 weeks after birth. Pups infected with hookworm larvae in their dams' milk typically begin to pass eggs in their feces 2 weeks after birth. It is important that pups receive treatment early in life if infection with roundworms or hookworms is suspected. To prevent peracute hookworm disease in unweaned pups of bitches harboring large numbers of somatic larvae, it might be necessary to treat the pups before hookworm eggs are detectable in fecal examinations. Canine en- doparasites are reviewed in Chapter 5 and discussed fully elsewhere (Georgi and Georgi, 1992~. SOCIALIZATION OF PUPS There is ample evidence of the importance of adequate socialization for the normal behavioral development of dogs (Clarke et al., 1951; Fox, 1968; Freedman et al., 1961; Houpt, 1991; Scott and Fuller, 1965~. The term socialization is somewhat confusing because it has been used to describe events, processes, and procedures. In the narrowest sense, socialization is the development of the primary social attachments that form between a pup, its dam, and its littermates during a critical or sensitive period in its behav- ioral development (Scott, 1968~. The process is not peculiar to dogs but occurs in many species of social mammals (see, for example, Cairns, 1966; Harlow and Harlow, 1969~. In a broader sense, socialization is the process by which pups form attachments to other dogs, people, and environments. Attachment formation might require nothing more than sufficient exposure to or experience with other dogs, people, and elements of the environment, which results in familiarity with a variety of stimuli (Cairns, 1966; Scott, 1963~. Breeds and individual pups differ in ease of socialization (Scott, 1970~. In any case, adequate socialization allows a pup to develop normal social relationships with other dogs and to adapt to pair or group housing, to adjust more easily to unfamiliar stimuli and environmental changes, and to accept handling with little or no fear and distress (Scott, 1980~. Sensitive Period for Socialization There is a sensitive period for socialization during which attachments form most readily and rapidly (Scott and Fuller, 1965~. The beginning of the period is marked by the startle response to sound at the age of approxi- mately 3 weeks. Also at 3 weeks, a pup begins to display distress vocaliza

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MANAGEMENT OF BREEDING COLONIES 45 lions when separated from its dam. Distress vocalizations are distinct from those made in response to fear (Davis et al., 1977), hunger (Compton and Scott, 1971; Scott and Bronson, 1964), or physical discomfort (Gurski et al., 1980~. Separation distress is greater in an unfamiliar pen (Elliot and Scott, 1961~. To minimize separation distress, pups should remain with their dams for at least their first 6 weeks. Ease of attachment formation varies between breeds and individuals but generally peaks between the age of 6-8 weeks (Scott and Bronson, 1964~. Although socialization probably occurs at a low rate throughout life, the end of the sensitive period is marked by the pup's increasing fear of the unfamiliar at the age of 12-14 weeks (Scott, 1962~. Consequences of Inadequate Socialization Pups that are inadequately socialized during the sensitive period exhibit abnormal behaviors, called kennel-dog or isolation syndromes, that are char- acterized by one or more of the following behaviors: generalized fearful- ness, fear-motivated aggression, timidity, immobility, or hyperactivity (Scott et al., 19674. Dogs that, as a result of inadequate socialization, become highly distressed when subjected to common laboratory Procedures (e.~. ~ , %, , . ~ By, ~ ~ ~ ~ ~ ~ . . ~ ~ en handling, walking on a leash, restraint, venipuncture, moves to different enclosures, and contact with other dogs) probably do not make good re- search subjects and might be in a compromised state of well-being. It has been reported that physiologic measurements on such dogs can fall outside normal limits (Vanderlip et al., 1 985b). Socialization Programs Providing contact and handling only during routine husbandry proce- dures might not be sufficient to produce behaviorally normal, cooperative research animals (Vanderlip et al., 1985a,b). Specific programs that ad- dress each aspect of socialization to dogs, to people, and to the environ- ment should be implemented. Programs that can be used as examples for providing adequate socialization have been reported (Vanderlip et al., 1985a,b; Wolfle, 1990~. The following are examples of elements that might be included in so- cialization programs: positive contacts with more than one person, opportu- nities to follow handlers, introduction to some type of restraint (e.g., a collar and leash), contacts with conspecifics other than littermates, and op- portunities to explore outside the kennel. Exploration might include expo- sure to floors of different textures, to a room with different lighting, to stairs, and to such equipment as exam tables, clippers, and scales. Expo- sures to those elements should be gradual and paired with positive reinforc

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46 DOGS: LABORATORY ANIMAL MANAGEMENT ers, such as food, petting, or verbal praise. Negative reinforcement and physical punishment can elicit aggressive or fearful behaviors and will make pups more difficult to handle. It is not necessary, or practical, to introduce pups to every type of environment, person, or animal to which they will later be exposed in order to provide adequate socialization. Evidence sug- gests that experience in coping successfully with change facilitates later success (Scott, 1980~. Thus, the adequacy of any socialization program can be determined by the ability of pups to adapt successfully to environmental changes with minimal behavioral and physiologic disruption. RECORD KEEPING Records on colony reproduction are essential. Individual records should contain the following minimal information on each bitch: . start date of each proestrus; dates of mating and stud dog's identification number; date on which bitch's diet should be increased (day 42 of gestation), date to move bitch to whelping facility (day 50), and range of expected whelping dates; actual whelping date, whelping complications, number and sex of live pups, number of stillbirths, and any obvious abnormalities in the pups; date to start weaning, bitch's distemper antibody titer (if known), and dates to deworm and vaccinate litter; and . daters) and details of disposition of litter. In addition, missed cycles, abortions, or any abnormal maternal behavior should be recorded. To facilitate review of the reproduction records of an entire colony, it is helpful to have a separate computerized or manual-entry spreadsheet that displays every reproductive cycle of each bitch in the colony. The spread- sheet is most useful if it lists the following information, organized chrono- logically by date of proestrus: . identification number of each bitch whose proestrus was first ob- served on that date; for each bitch bred, identification number of stud dog, first and last dates of mating, total number of matings, and calculated or expected dates for medical examinations, moving to whelping facility, and whelping; and expected date of next cycle. The spreadsheet should be updated periodically to include for each bitch the actual whelping date; the length of gestation; litter information, as described

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MANAGEMENT OF BREEDING COLONIES 47 above; the actual date of the next cycle; and the calculated interestrus inter- val. A computerized list can be sorted to review the breeding records of individual bitches and males over several years. Such a list also allows examination for trends in low fertility, long or short gestation lengths as indicators of poorly timed inseminations, number of matings per cycle, pro- jected periods during which several bitches will be in heat at the same time or no bitches will be in heat, and other matters that could reflect husbandry, management, or staff problems that need correction. REFERENCES AAFCO (Association of American Feed Control Officials), Canine Nutrition Expert Subcom- mittee, Pet Food Committee. 1993. AAFCO nutrient profiles for dog foods. Pp. 92-99 in Official Publication 1993. Atlanta: Association of American Feed Control Officials. Available from Charles P. Frank; AAFCO Treasurer; c/o Georgia Department of Agricul- ture; Plant Food, Feed, and Grain Division; Capitol Square, Atlanta, GA 30334. Amann, R. 1986. Reproductive physiology and endocrinology of the dog. Pp. 532-538 in Current Therapy in Theriogenology 2. Diagnosis, Treatment and Prevention of Repro- ductive Diseases in Small and Large Animals, D. A. Morrow, ed. Philadelphia: W. B. Saunders. Andersen, A. C. 1970. Reproduction. Pp. 31-39 in The Beagle as an Experimental Dog. Ames: Iowa State University Press. Andersen, A. C., and M. E. Simpson. 1973. The Ovary and Reproductive Cycle of the Dog (Beagle). Los Altos, Calif.: Geron-X. 290 pp. Bouchard, G. F., N. Solorzano, P. W. Concannon, R. S. Youngquist, and C. J. Bierschwal. 1991a. Determination of ovulation time in bitches based on teasing, vaginal cytology, and ELISA for progesterone. Theriogenology 35:603-611. Bouchard, G., R. S. Youngquist, B. Clark, P. W. Concannon, and W. F. Braun. l991b. Estrus induction in the bitch using a combination diethylstilbestrol and FSH-P. Theriogenology 36:51-65. Bouchard, G., H. Plata-Madrid, R. S. Youngquist, G. M. Buening, V. K. Ganjam, G. F. Krause, G. K. Allen, and A. L. Paine. 1992. Absorption of an alternate source of immunoglobu- lin in pups. Am. J. Vet. Res. 53:230-233. Breazile, J. E. 1978. Neurologic and behavioral development in the puppy. Vet. Clin. North Am. 8:31-45. Burke, T. J., ed. 1986. Small Animal Reproduction and Infertility. Philadelphia: Lea & Febiger. 408 pp. Cairns, R. B. 1966. Attachment behavior in mammals. Psychol. Rev. 73:409-429. Carmichael, L. E. 1983. Immunization strategies in puppies Why failures? Compend. Contin. Educ. Pract. Vet. 5:1043-1051. Christiansen, I. J. 1984. Reproduction in the Dog and Cat. London: Balliere Tindall. 309 PP Clarke, R. S., W. Heron, M. L. Fetherstonhaugh, D. G. Forgays, and D. O. Hebb. 1951. Individual differences in dogs: Preliminary report on the effects of early experience. Can. J. Psychol. 5:150-156. Compton, J. M., and J. P. Scott. 1971. Allelomimetic behavior system: Distress vocalization and social facilitation of feeding in Telomian dogs. J. Psychol. 78:165-179. Concannon, P. W. 1991. Reproduction in the dog and cat. Pp. 517-554 in Reproduction in Domestic Animals, 4th ea., P. T. Cupps, ed. New York: Academic Press.

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