1174 



HORMONAL REGULATION 



cognized. Their role in the expression of 

 sexual behavior in man is equivocal 

 (see chapter by Money; in addition, Cree\y 

 and Rea, 1940; Tauber, 1940; Carmichael, 

 Noonan and Kenyon, 1941 ; Filler and 

 Drezner, 1944; Carter, Cohen and Shorr, 

 1947; Kinsey, Pomeroy and Martin, 1948; 

 Kinsey, Pomeroy, Martin and Gebhard, 

 1953; Perloff, 1949; Waxenberg, Drellich 

 and Sutherland, 1959; and many others), 

 but for most vertebrates for which data 

 exist the full display of mating behavior is 

 given only when the gonadal hormones are 

 present. The relationship may be more di- 

 rect than any other between endocrine sub- 

 stances and behavior.^ When the gonads 

 are removed mating behavior becomes 

 greatly reduced in intensity or is no longer 

 displayed; on replacement of the hormone 

 it is again exhibited. If an inadequate 

 amount is given, a partial response may be 

 detected, or, as in a number of rodents and 

 the cow, when an improper balance is ad- 

 ministered an atypical pattern may be 

 shown (Boling, Young and Dempsey, 1938; 

 Melampy, Emmerson, Rakes, Hanka and 

 Eness, 1957). However, when the threshold 

 of the proper hormones is reached and, in 

 species requiring estrogen and progesterone, 

 the correct balance is given in the proper 

 sequence, the pattern of behavior identical 

 with that displayed before gonadectomy 

 is restored. In what follows, the observa- 

 tions and experiments supporting this con- 

 clusion are reviewed, factors which in- 

 fluence, modify, and limit the action of 

 gonadal hormones are enumerated, and sug- 



- From the fact that deviations of behavior are 

 well known in dysfunction of the parathyroid, the 

 thyroid, the pituitary, and the adrenal cortex, 

 and in hyperinsulinism (Hoskins, 1934; Shock, 

 11)44; Sherman, 1945; Medvei, 1949; Young, 1954), 

 it might be concluded that general behavior is de- 

 pendent on a normal functioning of these glands 

 and therefore that a direct relationship exists. 

 This may be true for cortisone and related steroids 

 secreted by the adrenal cortex (Rome and Brace- 

 land, 1950, 1951, 1952; Derbes and Weiss, 1951). 

 For the other endocrine organs, however, there 

 still is no uneqivocal proof of a one-to-one re- 

 lationship between the active principles produced 

 by these glands and the changes in behavior 

 associated with deficiencies or with excessive 

 quantities. An indirect relationship traceable to 

 derangements of metabolism or to physical de- 

 formity and handicap seems more likely. 



gestions are made with respect to the char- 

 acter and scope of problems awaiting fur- 

 ther study. 



The discussion will begin with a descrip- 

 tion of mating behavior, for the end points 

 investigators have used in obtaining their 

 data have been derived from the elements 

 or measures composing this behavior.^' * As 

 the substance of our description becomes 

 apparent, many will feel that a dispropor- 

 tional space has been given to the rat and 

 guinea pig. This is explained by the circum- 

 stance that the rat and guinea pig have 



^ Many articles in which mating behavior is 

 described are cited in the older reviews (Stone, 

 1932b, 1939b; Young, 1941 ; Beach, 1942e, 1946, 1947, 

 1948, 1951, 1952; Bullough, 1945; Hartman, 1945; 

 CoUias, 1950). A number of newer studies con- 

 taining bibliographies or unusually complete de- 

 scriptive accounts of mating behavior are noted 

 here (Schlosberg, Duncan and Daitch, 1949; 

 Baerends, Brouwer and Waterbolk, 1955; Morris, 

 1955b; Aronson, 1957; for fishes. Noble and Aron- 

 son, 1942; Aronson, 1944; Russell, 1955; for am- 

 phibians. Domm and Davis, 1948; Guhl, 1949; 

 Richdale, 1951 ; Hinde, 1952, 1955a, b ; Hale, 1955 ; 

 Morris, 1954; Wood-Gush, 1954; Moynihan and 

 Hall 1955; for birds. Macirone and Walton, 1938; 

 Stone and Ferguson, 1940; Snell, Fokete, Hummel 

 and Law, 1940; Pearson, 1940; Beach and Holz, 

 1946; Reed, 1946; Reed and Reed, 1946; Beach and 

 Pauker, 1949; Young and Grunt, 1951; Enders, 

 1952; Rowley and Mollison, 1955; Scott and 

 Lloyd-Jacob, 1955; Larsson, 1956; Green, Clemente 

 and de Groot, 1957; Rosenblatt and Aronson, 

 1958a, b; Beach and Rabedeau, 1959; for small 

 mammals including the rabbit, cat and dog. Bur- 

 ger, 1952; for the domestic pig. Hafez, 1952; for 

 the ewe. Carpenter, 1942a, b; Yerkes, 1943; Stellar, 

 1960; for infrahuman primates). 



*It is of historic interest that endocrinologists, 

 physiologists, biochemists, and anatomists who 

 have contributed so importantly to our knowledge 

 of the microscopic anatomy and physiology of 

 reproduction, have given little attention to prob- 

 lems centering around the hormonal regulation of 

 reproductive behavior. Much of the work in this 

 field has been done by psychologists and much of 

 the support that has encouraged effort in this 

 direction can be attributed to their foresight, 

 activity, and persistence. Endocrinologists, physi- 

 ologists, and anatomists have done little with 

 behavior, probably because of the former rather 

 generally held belief, best expressed perhaps by 

 Stockard and Papanicolaou (1919) and Moore and 

 Gallagher (1930), that behavioral end points are 

 so vague as to be of little value in experimental 

 studies. This circumstance in the development 

 of the subject is reflected by the absence of any 

 discussion of the hormones and behavior in most 

 textbooks of endocrinology. 



