HORMONES AND MATING BEHAVIOR 



1187 



cent additions to the information collected at 

 the time of estrus (Goy and Young, 1957b). 

 When the rat has been used there has been 

 a numerical evaluation of the responses such 

 as lordosis, quivering of the ears, darting, 

 and crouching when the female is ap- 

 proached by the male or fingered (Ball, 

 1937b), or the calculation of a quotient, 

 the copulatory quotient, which is the num- 

 ber of lordosis responses divided by the 

 number of times the female was mounted 

 by the male multiplied by 100 (Beach, 

 1944c). In a study of the female rabbit the 

 nvmiber of successful matings was divided 

 by the attempts made by the male. A pro- 

 portion was obtained which could be plotted 

 for each day (Klein, 1952). Receptivity of 

 the female cat can be assessed by daily 10- 

 minute mating tests with especially trained 

 males. The end point chosen for quantitative 

 assessment of the behavioral change was the 

 first occurrence of full mating accompanied 

 by the presence of sperm in the vaginal 

 smear (Michael and Scott, 1957) . 



Mounting, or pseudomale behavior, as 

 Morris (1955a) calls it, is measured in the 

 guinea pig by watching animals known to 

 be about to come into heat and counting 

 the number of times individuals mount other 

 animals (Young and Rundlett, 1939; Goy 

 and Young, 1957b) , and in the rat by divid- 

 ing the sum of points obtained in a test by 

 the duration of the test (Koster, 1943). 



Running activity, the first of the elements 

 of female sexual behavior to be studied 

 (Slonaker, 1912), has long been measured 

 by means of activity wheels. That developed 

 by Farris (1941) is probably the most elab- 

 orate. The number of revolutions of a turn- 

 table is registered by a magnetic counter 

 which is photographed at selected intervals 

 by a time-lapse mechanism. Total activity 

 including restlessness is better measured by 

 stabilimeter cages in which any movement 

 of the animal rocks the cage in a motion 

 which is recorded (Richter, 1927; Wilbur, 

 1936; Hunt and Schlosberg, 1939; Smith, 

 1940; Bousfield and Mote, 1943; Eayrs, 

 1954). Eayrs pointed out that, although 

 both types of apparatus are believed to 

 measure the same thing, the two techniques 

 measure components of activity having dif- 

 ferent motivational siiinificanco. 



The temporal relationships between lor- 

 dosis, male-like mounting, and running ac- 

 tivity are generally close because in lower 

 mammals all are displayed near the time of 

 ovulation. The quantitative relationships 

 are not close. Some rats are relatively in- 

 active at the time of estrus (Hitchcock, 

 1925; Stone and Barker, 1934). Male-like 

 mounting in guinea pigs is variable (Av- 

 ery, 1925), but the amount is not related 

 to the duration of estrus (Young, Dempsey 

 and Myers, 1935; Young, Dempsey, Hag- 

 quist and Boling, 1939). The variability 

 within each type of behavior has the effect 

 of creating differences between individual 

 females that are fully as striking as those 

 between males (Hitchcock, 1925, Young and 

 Fish, 1945, for running activity in the fe- 

 male rat; Young, Dempsey and Myers, 1935, 

 Young, Dempsey, Hagquist and Boling, 

 1939, for the length of heat and the amount 

 of mounting activity in the guinea pig; 

 Blandau, Boling and Young, 1941, for the 

 length of heat in the rat; Young and Orbi- 

 son, 1944, for the character of sexual re- 

 sponses in the chimpanzee; de Alba and 

 Asdell, 1946, for the strength of heat in the 

 cow; JMichael, 1958, for the behavior pat- 

 terns in the female cat). 



The length of heat in the guinea pig is 

 not related to the number of developing fol- 

 licles (Young, Myers and Dempsey, 1933) 

 or to the ovarian condition, at least, within 

 the limits of the ovarian pathology en- 

 countered by Young, Dempsey, Myers and 

 Hagquist (1938). Nor is the estrone content 

 of the follicular fluid necessarily related to 

 the degree of sexual desire in the mare (An- 

 drews and McKenzie, 1941). In what ap- 

 pears to be the single study in which such 

 data were collected, male-like mounting 

 was generally proportional to the number 

 of developing follicles (Young, Dempsey, 

 Myers and Hagquist, 1938), although nu- 

 merous exceptions were found. Cystic fol- 

 licles frequently occur in nvmphomanic 

 cattle (Pearl and Surface, 1915; Calder, 

 1927; Hammond, 1927; Fernandez, 1940; 

 Walton, Edwards and Hammond, 1940), 

 although not all cows with cystic ovaries 

 are nymphomanic (Casida, ]\IcShan and 

 Meyer, 1944). 



The absence of estrous behavior follow- 



