Berkeley Field Station for Behavioral Research Dog Studies
Dr. Frank A. Beach helped found the Field Station for Behavioral Research at University of California, Berkeley in 1963. Over the next decade, he and many other researchers, including Dr. Ian Dunbar, studied the sexual and maternal behavior of a colony of beagles. This research formed much of our early understanding of how dogs breed, before the rise of theriogenology as a field.
You may find that you know everything in these papers so consider this a history lesson that shows how little we knew about dogs as little as 40 years ago. However, I believe there are some important insights in this research for today’s breeders, such as the importance of amniotic fluid for dam-pup bonding. Here are just a few of the publications that came from their research. We have collected the highlighted articles or abstracts in PDFs for you that are linked to this page. Most of the full articles are available for purchase online if you want to dig in more deeply.
Enjoy!
Table of Contents
Coital Behaviour In Dogs Series
Beach, FA and BJ LeBoeuf (1967), Coital behaviour in dogs. I. Preferential mating in the bitch.
Animal Behaviour, vol 15, iss 4 (Oct), pp 546-548.
Abstract
Five male and five female beagles were raised together from puppyhood in a large field and were tested for copulatory behaviour when the females came into oestrus. Mating tests were repeated 8 to 13 months later during a second oestrous period. Females exhibited clear-cut preferences for particular males as sexual partners. Feminine rejection behaviour ranged from simple avoidance to active attack. Some females were more selective than others, but all showed discriminatory responses. Some males were rarely rejected by any bitch, whereas others were generally unpopular. It is suggested that any concept of sexual receptivity as an endogenously controlled condition leading to indiscriminated acceptance of all conspecific masculine partners must be evaluated separately for each species. The absence of preferential responsiveness on the part of the females should not be assumed a priori. Its existence or nonexistence can be established only by direct investigation.
Beach, FA (1968), Coital Behaviour in Dogs. III. Effects of Early Isolation on Mating in Males. Behaviour, Vol. 30, No. 2/3, pp. 218-238
Abstract
Three groups of male beagles were reared from weaning to sexual maturity under special circumstances and their responses to estrous females were compared in a series of mating tests. Five dogs were reared under conditions of semi-isolation (SI). They lived in individual cages and had very little physical contact with other animals throughout the experiment. The 5 males constituting the control group also inhabited individual cages but for 15 min. each day they were set free in the colony room where they could interact with other uncaged dogs. A final group of 5 males was reared together with 5 females in a 1-acre field. None of the subjects had contact with estrous bitches except during the mating tests. The copulatory performance of the control and the group-reared males was essentially the same, but that of the semi-isolates was deficient in one particular. SI males mounted estrous females as readily and as frequently as members of the other 2 groups, but the orientation of mounting was abnormal. Ninety-seven per cent of the mounts by control and group-reared males were oriented to the female’s rear, whereas 39 per cent of the mounts by SI dogs were directed to the head, side, or flank of the receptive bitch. One consequence of this difference was that SI males achieved intromission in only 24 per cent of their tests, as compared with 58 per cent and 54 per cent for the control and group-reared animals.
Beach, FA (1970), Coital Behaviour in Dogs. VIII. Social Affinity, Dominance and Sexual Preference in the Bitch. Behaviour, Vol. 36, No. ½, pp. 131-148
Abstract
Six male and 5 female dogs were reared together in an outdoor compound (20.5 m2) from the time they were 12-14 weeks old until the females had their first estrous period at 28-36 weeks of age. During this time all animals were observed in 3 tests designed to measure social affinity or attraction between males and females, a test for social dominance and a test which revealed each female’s tendency to accept or reject each of the males in copulation.
In Free Play (FP) Tests all 11 dogs were released in the living area and records were made of the frequency of social interaction between every female and every male. Dyadic Roving (DR) Tests involved observations of one male and one female at a time while they were free to move about the living area in the absence of other dogs. The frequency and amount of time each female interacted with every male were recorded. In the Tethered Male (TM) Tests one male was chained to a stake in the center of the living area and females were released in the area one at time. Notes were made of the amount of time each female spent “visiting” each male.
Results of these 3 tests of social affinity showed that females differed from one another with respect to their tendency to interact with any male, some bitches being much more “social” than others. Each female displayed a pattern of preferences, spending more time with some males than with others. There was some tendency for females to respond more frequently to their brothers than to unrelated males. Comparing the results of the 3 tests revealed a definite though fairly low degree of inter-test consistency in the social preferences of the 5 females.
During the Bone Possession Tests one male and one female were given simultaneous access to a large bone and records were made of the amount of time each dog maintained possession. On the average males tended to surpass females but there were marked individual differences in both groups. Lolita, the most dominant female, controlled the bone an average of 52.17% of the time in her tests with the 6 males, whereas the comparable score for Kathy was 13.00%. Among the males Cassius was in possession of the bone an average of 90.80% of the test time while Don controlled the incentive only 58.80% of the time. It was noted that the most dominant male and female were individuals who ranked low in frequency of social interactions as measured by the DR and TM tests. Success in maintaining possession of the bone was unrelated to physical size as measured by body weight.
Mating Tests conducted when 4 of the females were in estrus revealed the existence of clear-cut sexual preferences on the part of each bitch. Patterns of choice varied from one female to the next.The same male might rank first on the preference scale of one bitch and last on that of another. Comparisons between the degree to which each female accepted each male as a sexual partner and the extent to which that same pair had interacted in the tests for social affinity indicated no relationship whatsoever. Similarly the results of the dominance tests bore no apparent relation to patterns of feminine sexual preference revealed by the mating tests. It was concluded that (1) anestrous females exhibit varying degrees of social affinity for different males with whom they have been reared, but (2) under the influence of hormonal conditions associated with estrus these social preferences are obliterated or overridden and an entirely new pattern of heterosexual affinities and aversions emerges.
Beach, F. A. (1970). Coital behavior in dogs. IX. Sequelae to “coitus interruptus” in males and females. Physiology & Behavior, 5(3), 263-268.
Abstract
During normal mating male and female dogs form a genital lock during which the penis is so swollen within the vagina that the pair cannot separate until detumescence begins. After a normal lock (15 min +) most males and females are relatively calm and inactive. In some case a lock is broken within a few minutes after insertion and when this occurs the subsequent behavior of both sexes may undergo a sudden change. Most females become exceedingly hyperactive and sexually aggressive toward the male. In the same circumstances some males exhibit a phantom lock. For 5–15 min they stand nearly motionless in the posture assumed during a normal lock. The back is arched, tail fully erect and rhythmic ejaculatory contractions occur. Under these conditions males show little or no response to exteroceptive stimulation. The neurophysiological basis for the reactions of both sexes is discussed.
Beach, F. A., & Kuehn, R. E. (1970). Coital behavior in dogs. X. Effects of androgenic stimulation during development on feminine mating responses in females and males. Hormones and Behavior, 1(4), 347-367.
Abstract
Female beagles were exposed to androgenic stimulation prenatally, neonatally, or both. A fourth group was ovariohysterectomized in adulthood and never given androgen. Male beagles were castrated at birth, just before puberty, or in adulthood. All males and females were injected with estrogen and progesterone during maturity and tested with experienced stud males for the display of sexually receptive behavior. Females spayed as adults exhibited the most complete and intense mating responses and were the most stimulating to stud males. Females exposed to androgen before but not after birth ranked second in both of these respects. Females treated with androgen during infancy were much less receptive than those treated prenatally but exhibited some responsiveness to ovarian hormones. Females receiving androgen both pre- and postnatally did not differ from males castrated in adulthood. Neither group showed any clear-cut behavioral response to estrogen and progesterone. Neonatally and prepuberally castrated males were noticeably influenced by the hormones on some behavioral measures, with the former group being somewhat more responsive than the latter.
Beach, F. A., Kuehn, R. E., Sprague, R. H., & Anisko, J. J. (1972). Coital behavior in dogs. XI. Effects of androgenic stimulation during development on masculine mating responses in females. Hormones and Behavior, 3(2), 143-168.
Abstract
Six groups of adult beagles were tested for the display of masculine mating responses to sexually receptive bitches. Treatment and constitution of the several groups were as follows: females whose mothers were injected with TP for 10 days during the second trimester, females implanted with a testosterone pellet 1–3 days postpartum, females whose mothers were injected with TP during pregnancy and who received a testosterone pellet 1–3 days postpartum, females ovariohysterectomized in adulthood but given no androgen during development, males castrated within 48 hr postpartum, and males castrated in adulthood. All groups received periodic injections of TP as adults before and during mating tests.
Prenatal androgenic stimulation prevented development of an external vagina and induced development of a penis in all cases. Treatment starting at birth resulted in clitoral hypertrophy but the vagina was patent.
Females untreated during development and ovariectomized as adults showed relatively little masculine behavior when injected with TP in adulthood. Females untreated before birth but given testosterone implants during the neonatal period were not appreciably “masculinized.” In contrast, females treated in utero but receiving no treatment in infancy displayed significantly more masculine sexual activity than either of the two preceding groups. The most frequent and complete masculine behavior shown by any of the females was exhibited by those which had been exposed to androgen both before and immediately after birth.
Most of the males castrated in adulthood and injected with TP 15–19 months later displayed the full complement of coital responses characteristic of the normal male. Males castrated at birth and given TP as adults showed much more masculine sexual behavior than any group of females, but on most measures were noticeably inferior to male subjects castrated after attainment of sexual maturity.
Male-Female Attraction and Responses
Beach, FA (1974), Effects of gonadal hormones on urinary behavior in dogs. Physiology & Behavior, vol 12, Iss 6 (June): pp 1005–1013.
Abstract
The effects of testicular and ovarian hormones on urinary frequency and posture were studied in 8 groups of dogs. Three of the 4 female groups had been exposed to androgenic stimulation in utero, Berkeley Field Station for Behavioral Research Dog Studies 8 | Page © Avidog International LLC 2013 neonatally, or both in utero and in infancy. The fourth group of females consisted of ovariectomized controls. Males were intact or castrated as adults, as juveniles, or as neonates.
Frequency of urination was increased in all groups by estrogen and by testosterone. Urinary posture was unaffected in males by castration in the adult or juvenile stages. Neonatal castration of males resulted in periodic regression from the adult male posture to the immature male posture.
Control and prenatally androgenized females urinated in the feminine position. Females treated
with large amounts of testosterone in infancy showed a limited degree of masculinization of urinary posture. Females exposed to testosterone before and immediately after birth urinated as females about 50% of the time and as adult males for approximately half of their urinations. Exogenous estrogen or androgen administered in adulthood had no effect on posture assumed for urination.
Beach, F. A., Buehler, M. G., & Dunbar, I. F. (1983). Development of attraction to estrous females
in male dogs. Physiology & behavior, 31(3), 293-297.
Abstract
In simultaneous choice tests male beagles were allowed to visit a caged female in estrus, or caged, spayed female not in estrus. Males were tested periodically from 1–3 to 22–24 months of age. The 3 subject groups were normal males (Group N), males castrated 4–7 days postpartum and injected with testosterone propionate (TP) until they were 3 months old (Group TPTC), and males castrated 4–7 days but given no hormone treatment, i.e., untreated castrates (Group UC). A statistically reliable preference for visiting the estrous rather than the nonestrous female first appeared in N males at 4–6 months, in TPTC males at 1–3 months, and in UC males at 10–12 months. In N males attraction to the estrous female (measured by time spent visiting her) increased progressively from 4–6 to 16–18 months. In the same period concentration of plasma testosterone rapidly increased, reached a peak at 10–12 months, and then declined. UC males exhibited no significant increase in attraction to the estrous female from 10–12 months (when a preference first appeared) to 22–24 months. They then received 10 injections of TP after which their visiting time to the estrous female was equal to that of N males. TPTC males exhibited a precocious preference for the estrous female at 1–3 months, while they were receiving TP, and there was no decrease in strength of attraction in the following 9 months during which time no hormone was administered. Between 10–12 and 13–15 months, still without exogenous androgen, visiting to the estrous female began to increase and continued to do so until 19–21 months.
Doty, R. L., & Dunbar, I. (1974). Attraction of beagles to conspecific urine, vaginal and anal sac secretion odors. Physiology & Behavior, 12(5), 825-833.
Abstract
Attraction of male and female Beagles to conspecific urine, vaginal and anal sac secretion odors was examined in four experiments. Males spent more relative time investigating female urine odors than odors of vaginal or anal sac secretions. Sexually experienced males, but not sexually inexperienced ones, spent more time investigating estrous than diestrous female urine and vaginal odors. Anal sac secretions from estrous bitches were not more attractive to males than those from diestrous bitches. Estrous females spent no more time than diestrous ones in the investigation of male anal sac secretion and urine odors. Male urine and anal sac secretions elicited little investigation from male conspecifics. Females spent more time investigating female urine odors than female anal sac or vaginal secretion odors, and exhibited a slight general preference for diestrous over estrous stimuli. A positive correlation between the odor investigation times of this study and investigation times of comparable animals to conspecifics in a social situation suggests odor preferences are relatively good indicators of social preferences, and vice versa, in this breed.
Dunbar, I, MG Buehler, FA Beach (1980), Developmental and activational effects of sex hormones on the attractiveness of dog urine. Physiology & Behavior, Vol. 24, Iss. 2 (Feb): pp 201-2014.
Abstract
Adult male dogs were exposed to urine from five types of experimental donors: females ovariohysterectomized as adults (FOA), females treated with testosterone in utero (FTU), females treated with testosterone in early infancy (FTI), females treated with testosterone in utero and in early infancy (FTUI), males castrated as adults (MCA); and also to control urine from intact stud males (SM). Urine was collected from the five experimental groups of donors during three phases of the experiment: (a) during a period of no hormone treatment (NH); (b) following treatment with estradiol (E); and (c) following treatment with testosterone (T). Intact males were given no hormonal treatment and served as subjects and urine donors. Subjects spent about the same amounts of time investigating control SM urine in all three phases. In the NH phase, the investigation times for the five experimental samples were significantly greater than that of the SM sample. However, there were no significant differences between the investigation times of the five samples. During the E phase, in comparison with the NH phase, the investigation times were greater for all experimental samples. The increase in attractiveness was significant for FOA and FTU urine. During the T phase, compared with the NH phase, the investigation times were less for four experimental samples (FTU, FTI, FTUI and MCA urine), and FTI, FTUI and MCA urine were no longer preferred to the SM control. However, the decrease in attractiveness was significant for MCA urine only.
Dunbar, I. F. (1977). Olfactory preferences in dogs: the response of male and female beagles to
conspecific odors. Behavioral Biology, 20(4), 471-481.
Abstract
In Experiment 1, male dogs were observed visiting either a tethered male or a tethered nonestrous female. The duration and frequency of visits and the parts of the stimulus animal that were investigated were recorded. In Experiment 2, male and female dogs were observed in a series of two-choice tests to determine their visiting preferences toward males and nonestrous and estrous females. In Experiment 3, similar two-choice olfactory tests were employed to demonstrate investigatory preferences for urine, feces, anal gland and vaginal secretions, saliva, and ear wax samples from the stimulus animals. The females were brought into artificial estrus with injections of estradiol benzoate, and changes were observed in their visiting and olfactory preferences and in their attractiveness to male dogs. Males spent less time visiting males than they spent visiting females, regardless of the physiological condition of the latter. However, the visiting time devoted to estrous females was longer than that spent with females not in heat. These preferences may be partially explained by the findings that male dogs find estrous urine and vaginal secretions more attractive than their nonestrous counterparts and that they prefer female urine and feces compared to corresponding samples from male dogs. Following the induction of artificial estrus, female subjects spent significantly more time investigating male dogs and male urine when compared to female dogs and female urine, respectively. Male dogs and to a lesser extent bitches urinated more frequently in the vicinity of the preferred animals and odor stimuli.
Bitch Behavior
Beach, F. A., Dunbar, I. F., & Buehler, M. G. (1982). Sexual characteristics of female dogs during successive phases of the ovarian cycle. Hormones and Behavior, 16(4), 414-442.
Abstract
Five female beagles were examined periodically throughout their second biannual cycle of proestrus, estrus, metestrus, and anestrus for fluctuations in several physiological and behavioral charateristics. Physiological measures included plasma levels of estradiol and progesterone as well as the vulvar swelling and vaginal bleeding that begin with proestrus and continue in estrus.
Behavioral tests revealed changes in female receptivity, attractivity, and proceptivity. Estradiol increased prior to the onset of proestrus and declined during estrus. Progesterone levels were low during most of proestrus, increased from the beginning to the end of estrus, and remained elevated during the first month or more of metestrus. While they were in proestrous females were not receptive but they and their vaginal secretions were highly attractive to males. Attractivity remained high throughout estrus and declined abruptly in the next 24 hr. Receptivity increased over the first 3 days of estrus and continued at a high level until the last 3 days during which it decreased slightly, and then dropped nearly to zero within the next 24 hr. “Sexual reflexes” of the vulva, tail, and hindquarters followed the same course of changes described for attractivity. Proceptive behavior, including seeking proximity to caged males and display of solicitation responses during mating tests was characteristic of females during proestrus and estrus but uncommon or absent in other phases of the cycle. It is hypothesized that in the natural cycle, attractivity and proceptivity develop during proestrus as a consequence of rapidly increasing secretion of estrogen. The onset of receptivity is due to synergistic action of estrogen and progesterone. Estrogen secreted during proestrus primes the system, and progesterone secreted just before and during estrus activates it. Termination of receptivity is thought to be due to the marked decline in estrogen, to inhibition by high concentrations of progesterone, or to a combination of these factors.
I Dunbar, E Ranson, M Buehler (1981), Pup retrieval and maternal attraction to canine amniotic
fluids, Behavioural Processes, Vol 6, Iss 3 (Oct), pp 249–260.
Abstract
Three purebred female beagles were observed with both their first and second litters. Dams were given three separate simultaneous-choice retrieval tests:
(1) male vs. female pup from their own litter;
(2) own pup vs. alien pup of the same sex and similar age;
(3) own pup daubed with amniotic fluid vs. own pup of the same sex treated with water.
In addition, the maternal response to amniotic fluids was observed when one pup in the litter was treated with amniotic fluids and all other pups were treated with water. There was no evidence to suggest that pups were retrieved preferentially on the basis of their sex. One female retrieved her own but not alien pups, whereas another female made no such discrimination and readily retrieved alien pups in addition to her own.
Retrieval behavior developed at the time of each whelping and normally lasted until the pups were about 5 days old, although a bitch would retrieve younger (alien) pups up to 14 days post partum. Maternal bitches were strongly attracted towards amniotic fluids: they investigated pups daubed with amniotic fluids to a significantly greater extent than control pups treated with water. The attraction of maternal females towards amniotic fluids developed at the time of each whelping and persisted for up to 30 days, well beyond the time that a whelping bitch would normally be exposed to her own fetal fluids. A possible role for amniotic fluids in the development of maternal behaviour and the establishment of the maternal/puppy bond is discussed.
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Breeding/Coital Behavior in Dogs Series
This eleven-part series of articles on sexual behavior in dogs is a mix of natural and manipulated
experiments on the beagles in the Berkeley colony. The series includes:
FA Beach, BJ LeBoeuf – Coital behaviour in dogs. I. Preferential mating in the bitch.
Animal Behaviour, 1967
Beach, F. A., Rogers, C. M., & LeBoeuf, B. J. (1968). Coital behavior in dogs II: Effects of estrogen on mounting by females. Journal of Comparative and Physiological Psychology, 66(2), 296.
Beach, F. A. (1968). Coital behavior in dogs. III. Effects of early isolation on mating in males. Behaviour, 218-238.
Beach, F. A., & Merari, A. (1968). Coital behavior in dogs. IV. Effects of progesterone in the
bitch. Proceedings of the National Academy of Sciences of the United States of America, 61(2), 442.
(Full article available here.)
Beach, F. A., & Merari, A. (1970). Coital behavior in dogs: V. Effects of estrogen and progesterone
on mating and other forms of social behavior in the bitch. Journal of Comparative and Physiological
Psychology; Journal of Comparative and Physiological Psychology, 70(1p2), 1.
Beach, F. A. (1970). Coital behavior in dogs: VI. Long-term effects of castration upon mating in the
male. Journal of Comparative and Physiological Psychology; Journal of Comparative and Physiological Psychology, 70(3p2), 1.
Beach, F. A. (1970). Coital behaviour in dogs. VIII. Social affinity, dominance and sexual preference
in the bitch. Behaviour, 131-148.
Beach, F. A. (1970). Coital behavior in dogs. IX. Sequelae to “coitus interruptus” in males and
females. Physiology & Behavior, 5(3), 263-268.
Beach, F. A., & Kuehn, R. E. (1970). Coital behavior in dogs. X. Effects of androgenic stimulation
during development on feminine mating responses in females and males. Hormones and
Behavior, 1(4), 347-367.
Beach, F. A., Kuehn, R. E., Sprague, R. H., & Anisko, J. J. (1972). Coital behavior in dogs. XI. Effects of androgenic stimulation during development on masculine mating responses in females. Hormones and behavior, 3(2), 143-168. Male-Female Attraction and Responses
Beach, FA (1974), Effects of gonadal hormones on urinary behavior in dogs. Physiology & Behavior, vol 12, Iss 6 (June): pp 1005–1013.
Beach, F. A., Buehler, M. G., & Dunbar, I. F. (1983). Development of attraction to estrous females in male dogs. Physiology & behavior, 31(3), 293-297
Doty, R. L., & Dunbar, I. (1974). Attraction of beagles to conspecific urine, vaginal and anal sac secretion odors. Physiology & behavior, 12(5), 825-833.
Dunbar, I., Buehler, M., & Beach, F. A. (1980). Developmental and activational effects of sex
hormones on the attractiveness of dog urine. Physiology & Behavior, 24(2), 201-204.
Dunbar, I. F. (1977). Olfactory preferences in dogs: the response of male and female beagles to conspecific odors. Behavioral Biology, 20(4), 471-481.
Bitch Behavior
Beach, F. A., Dunbar, I. F., & Buehler, M. G. (1982). Sexual characteristics of female dogs during
successive phases of the ovarian cycle. Hormones and Behavior, 16(4), 414-442.
Dunbar, I, E Ranson, M Buehler (1981) Pup retrieval and maternal attraction to canine amniotic
fluids. Behavioural Processes, Vol 6, Iss 3 (Oct), pp 249–260
