Coital Behavior in Dogs
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Journal 0/ Comparative and Physiological Psychology 1968, Vol. 66, No.2, 296-307 COITAL BEHAVIOR IN DOGS: EFFECTS OF ESTROGEN ON MOUNTING BY FEMALESl FRANK A. BEACH, CHARLES M. ROGERS, University of California, Bej·keley Yerkes Regional Primate Research Center, A tlanta , Geoj·gia AND BURNEY J. LEBOEUF Unive1"Sity of California, Santa C7·UZ Female dogs mounted other females more frequently when the 2nd bitch was in estrus, but the hormonal state of the mounting individual had little influence upon mount frequency; estrogen rendered a female more pro vocative or stimulating and more cooperative or receptive to mounting by another dog. During heterosexual tests estrous females tended to mount males selectively, responding readily to some masculine partners and never mounting others; in limited tests anestrous females did not mount males. Estrogen facilitated expression of specific feminine copulatory reactions and increased tendencies to display various positive social responses to dogs of · either sex. Whether or not the hormone direcUy enhanced the female's tend ency to mount a conspecific partner was not determined. Female mammals of several species sexual receptivity (Young, Dempsey, Hag sometimes mount congeners and display quist, & Boling, 1939; Young & Rundlett, elements of the copulatory pattern of con 1939). specific males (Beach, 1967). Animal A study of female rats by Beach and husbandrymen report that female ungu Rasquin (1942) indicated that there might lates are most likely to mount while they be marked species differences in the hor are in estrus, although mounting by an monal control of mounting behavior. The estrous females has been observed. The incidence of mounting did not bear any implication that ovarian hormones may be systematic relation to the estrous cycle; involved is strengthened by a report that spayed females mounted as often as intact heifers implanted with pellets of stilbestrol individuals; and administration of estro mounted other animals so frequently and gen and progesterone to ovariectomized with such force that 20% of the treated fe animals did not increase the frequency males suffered pelvic fractures (Folley & with which they mounted, although it did Malpress, 1944). induce sexual receptivity. At the same time The first systematic studies of mounting it appeared that the hormonal condition in females were those of W. C. Young and of the stimulus female had a definite in his associates, who investigated the hor fluence upon the mounting responses of her monal control of this behavior in guinea partner. Females mounted other females pigs (Young, Dempsey, & Myers, 1935). In which were in estrus much more often and a series of investigations these workers more vigorously than they mounted dies amassed proof that for this species mount trous stimulus animals. ing is rare in diestrous females, is con The present report is an account of three centrated at the time of estrus, and can be experiments dealing with mounting be produced in spayed individuals by ad havior in female dogs. The results show ministration of the same combination of that this activity constitutes a complex estrogen and progesterone which induces form of social interaction which may be influenced by a combination of factors in 1 This research was supported in part by United cluding both the physiological state of the States Public Health Service Research Grants M-783 and MH-04000 from the National Institute mounting bitch and the stimulus qualities of Mental Health. of the potential mountee. 296 MOUNTING BY FEMALE DOGS 297 EXPERIMENT 1 Although the original purpose of these experi ments was to investigate the hormonal correlates This experiment, patterned after the of mounting, it soon became obvious that we were earlier study of rats by Beach and Rasquin not dealing with a simple reflexive response. In (1942), was designed to measure the ef stead, we were confronted with a complex form of interindividual behavior which could not mean fects of induced estrus upon homosexual ingfully be studied in isolation from several closely mounting in bitches which were also tested associated types of interaction. Mounting re for copulatory reactions to sexually active sponses appeared in a matrix of acts shown by one males. individual toward a conspecific partner and the entire pattern of social responses had to be ex NIethod amined if the presence or absence of mounting was to be understood. For this reason, TecOl·ds Subjects. Ten females and 11 males were pur taken during all observation periods included chased as adults from animal dealers. A few were notes on the frequency and timing of all forms of beagles but the majority were mongrels whose social activity. exact age and previous history were unknown. For the purpose of these investigations any be Three of the bitches were spayed and seven were havior was defined as "social" if it involved inter intact. action between the two Ss under observation. A H oll,sing and maintenance. From the time of "positive" social response was any act by one S their acquisition until the end of the experiment which was clearly oriented to the second S and Ss were housed in individual cages (30 X 31 X 39 which had· the effect of initiating or prolonging in.) in a basement room with forced ventilation nonantagonistic interaction between the members and artificial illumination. of the pair. "Negative" social acts included at Once each day for approximately 15 min., Ss tack, threat of attack, flight, and avoidance. These were released in small groups on the floor of the clearly were social in the sense of being oriented colony room while their cages were being cleaned, to another S, but they were classified as negative Except for these brief exercise periods no S had because they tended to prevent or to terminate physical contact with another except during the interaction. The positive social responses (PSR) behavior tests. regularly scored were as follows. (a) Rear mount: Testing and scoring procedures, All females S mounted the mountee from posterior position were observed with males during mating tests con with forelegs clasped about the flanks of the ducted while the bitch was in estrus; they also mountee; this is the coital position of the male were tested in homosexual pairs during estrus and during mating. (b) Rear mount with thrusts : anestrus. Estrus was induced by administration of S's hindquarters were moved rhythmically toward estradiol benzoate." Treatment consisted of two and away from the rump or perineal area of the intramuscular inj ections of .38 mg/kg spaced 48 moun tee; when the male copulates, this activity hr. apart. Females became receptive to the male precedes insertion. (c) Head mount : S approached 3-4 days after the second inj ection. Intact indi mountee from anterior position and grasps the viduals were injected during their natural period latter's neck or head between the forelegs; this of anestrus and their responses to hormone treat may have been accompanied by pelvic thrusting. ment did not differ in any detectable fashion from (d) Side mount: mountee was grasped, with or that of spayed females. without thrusting, from the side. (e) Investiga All tests were conducted in a basement room tion: S sniffed or licked the partner's body; the with a floor area of 120 sq. ft. In each hetero most frequent target of investigation was the sexual test the male was given a 5-min. period of anogenital area and the second was the head, adaptation before the estrous female was intro particularly the ears and mouth; investigation duced. Testing continued until a successful mating could be mutual and simultaneous. (f) Paws on occurred, or until 10-15 min. had elapsed and it body: S placed one or both forepaws on the part was apparent that the pair under observation was ner's body; frequently this appeared to be an not going to copulate on that day. In homosexual abortive attempt to mount, but in other instances tests one female was put in the observation room one dog merely lifted a forepaw and struck or 5 min. before introduction of the second S and the brushed against the head, shoulder, back, or rump pair was left together for 10 min. During all tests of the other. (g) Approach: S moved after a pe o watched Ss through a one-way vision window riod of separation to within one body length of and dictated into a recorder a running account the second dog. (h) Prance: S faced the head of of the behavior. the partner, flexed the forelegs so that its head was lower than its hindquarters, and either 2 The hormone preparation was generously sup "drummed" on the ground with a staccato move plied by Schering Corporation, Bloomfield, New ment of the forepaws or "bounced" up and down, Jersey. Subsequent experiments have indicated alternately elevating and lowering the entire body that doses considerably smaller than the one used while maintaining the "head-down-rump-up" pos in these studies might have been equally effective ture; in this position the "prancing" individual in inducing sexual receptivity. sometimes jumped laterally from left to right and 298 F. A. BEACH, C. M. ROGERS, AND B. J. LEBOEUF back again, but orientation to the face or head of females were in induced estrus, but two Ss the second dog was continuous; prancing com were later bred to males when they were in monly was accompanied by barking directed to ward the second animal. (i) Present: S, most often natural estrus and all mounting responses an estrous female, oriented her hindquarters to the were recorded. During tests in induced partner in such a way that her tail region was estrus one S mounted the male in 7% of 27 pointed directly toward the head of the "pre tests and the second S mounted in 5% of sentee"; this orienting movement frequently was 20 tests.