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Behavioral : An Emerging Discipline: How affect—and are affected by—behavior is a subject of increasing interest to students of human and animal behavior Author(s): Frank A. Beach Source: American Scientist, Vol. 63, No. 2 (March-April 1975), pp. 178-187 Published by: Sigma Xi, The Scientific Research Society Stable URL: http://www.jstor.org/stable/27845362 . Accessed: 21/05/2013 16:09

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This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions Franka. Beach Behavioral Endocrinology: An Emerging Discipline

How hormones affect?and are affected by? behavior is a subject of increasing interest to students of human and animal behavior

The first experiment in behavioral the concept of hormones as "chemi in covariation is the rule?which is endocrinology was performed in cal messengers" was proposed and to say the existence of correlations 1849 at the University of G?ttingen the science of endocrinology was involves variables other than just when Professor A. A. Berthold cas born. From its infancy, specialists the and the behavior trated cockerels and observed the in the new field maintained a genu under examination. For example, well-known changes always pro ine if peripheral interest in endo administration of small amounts of duced by such an operation, in crine effects on behavior; but for induces sexual receptivity cluding atrophy of the comb and half a century progress in this par in female but not in male rats; and spurs, cessation of crowing, and loss ticular aspect of endocrinology was stimulates male birds of sexual interest in hens. Bert slow, and only during the last fif to attack other males but not to hold's original contribution was the teen years has its development aggress against females. discovery that these postoperative been so dramatically accelerated changes could be prevented if he that a new discipline seems aborn The interactions and interdepen returned a single testis to the bird's ing. It is too early for general laws dencies which constitute the fourth body cavity where it would have no or comprehensive theories, but a problem area are numerous, com connections with the nervous sys few organizing principles seem to plex, and representative of the most tem but could establish an ade be crystallizing out of the accumu dynamic aspects of behavioral en quate blood supply. He concluded lating evidence, and research has docrinology. They reflect the fact that "the testes release something progressed far enough to show that many hormone-behavior rela into the blood that maintains male where the important problems lie tionships are reciprocal or linked in behavior and the secondary sex and what the principal methods are other ways. Stimuli aroused by be characters" (Turner and Bagnara by which they are likely to be havior sometimes feed back into 1971). solved. the brain and thence to the endo crine system, where they excite or The mysterious "something" was The major problems or problem inhibit the secretion of hormones, the hormone testosterone, but fifty areas discernible today can be clas which in turn exert additional ef years and more were to pass before sified in four interrelated categories fects on behavior. (Beach 1974). Covariation refers to correlations between endocrinologi Covariation Frank A. Beach is Professor of Psychology at cal and behavioral events. The first the University of California, Berkeley. After step in analyzing any correlation is Most of our information about co a Ph.D. in at the Uni obtaining psychology to determine its reliability or con variation of hormones and behavior versity of Chicago in 1940, he worked as As sistency; only after this is estab has come from three sources: di sistant Curator and Chairman of the De (1) partment of Animal Behavior at the Ameri lished is the problem of causality agnosis and treatment of endocrine can Museum of Natural History. He then examined. As soon as the reliability and behavorial pathology, (2) study 10 in the spent years Department of Psy of a correlation has been quanti of behavioral changes occurring in chology at Yale before going to University fied, the next task is to identify the concert with normal fluctuations in Berkeley in 1958. A member of the Ameri mechanisms the concentration of hor can Philosophical Society and the National mediating involved, specific Academy of Sciences, Dr. Beach has re thus assuring the existence of a mones, and (3) experimentation in ceived many awards for his research. His causal relationship and increasing the laboratory, clinic, and field. current work centers on sex in differences our understanding of how the hor the behavior animals and humans and of mone's behavioral are Glandular malfunction with their causes. This article is based on the consequences behav Kenneth Craik Memorial Lecture at St. brought about. ioral consequences is dramatically Johns College, Cambridge University. A illustrated: by the masculinization detailed version includes more, (Beach 1974) Research in behavioral endocrinolo of baby girls which occurs when de a comprehensive bibliography and detailed gy has progressed far enough to fective adrenal glands of the fetus documentation for general statements in show that instances of covariation secrete excessive amounts of male cluded here. Address: Department of Psy are chology, University of California, Berkeley, infinite but simple one-to-one hormone during pregnancy. The re CA 94720. relationships are rare. Contingency sulting pathological condition,

178 American Scientist, Volume 63

This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions known as congenital adrenal hyper female's behavior changes radically plasia or the adrenogenital syn as her hormonal tides ebb and flow. drome (AGS), includes modifica A woman's cycle, like that of most tion of the external but not the in other primates, is marked by regu ternal sex organs, so that some af lar periods of menstrual bleeding, flicted girls are born with a penis but otherwise it is quite similar to rather than a vagina. The genital the "estrous cycles" of lower mam anomaly can be surgically correct mals and tends to be accompanied ed, but some authorities believe by characteristic psychological or prenatal masculinization also af behavioral changes. Psychiatrists at fects the brain and thus influences the Stanford Medical School report sexually dimorphic behavior. that nearly one-third of American women experience some degree of Figure 1. An example of the normal varia emotional distress Drs. John Money, of Johns Hopkins tion in the endocrine system which is coinci (irritability, University Medical School, and dent with important behavioral changes is mood swings, depression, tension) testosterone in of Anke Ehrhardt, of the State Uni the increase in the blood just before or during menstruation, human males during adolescence (August, versity of New York, compared a when the is producing almost Grumbach, and Kaplan 1972). The colored number of AGS with normal no hormones at all and the individ girls segment denotes the critical range for the a girls the same age and found that onset of nocturnal emission, masturbation, ual is therefore in temporary state during childhood the former were dating, and first infatuation in the majority of withdrawal or deficiency of adolescent males "psychologically masculinized" ac (Ramsey 1943; Kephart (Hamburg, Moos, and Yalom 1973). cording to the following criteria 1968). (Money and Ehrhardt 1968): (1) From earliest infancy they pre Secretion of estrogen peaks at mid ferred boys' toys to dolls and other cycle, and this is the time when fe male hormone, but can be induced (2) they pre male animals are sexually attrac "girls' playthings," in most cases if testosterone treat ferred to play with boys and not tive, meaning that their stimulus ment is begun while the boy is still with other girls, (3) they were so value formales is high; sexually re in the adolescent age range. cially competitive in boys' groups, ceptive, meaning that they will co (4) they preferred boys' wearing ap operate in coition; and sexually In all mammals the secrete parel to frilly dresses, (5) they ex proceptive, meaning that they will estrogen and progesterone periodi pressed little or no interest in han assume initiative in seeking, solicit cally, although the periods may dling or caring for babies, (6) they ing, and stimulating males. Attrac come only once a year, as in sea rarely daydreamed of marriage and tivity and receptivity are familiar sonal breeders, or every four days, family but often of achievement concepts, but female proceptivity is as in the hamster. In all species the and careers, and (7) they consid often neglected, especially by male ered themselves and were consid investigators arid theorists. The ef ered by others to be tomboys. fect of ovarian hormones on male Money and Ehrhardt describe these seeking behavior in female dogs is manifestations of masculinization 35r shown in Figure 2, based on a study to be obvious but transitory, con Experimental females in which the roving bitch was free no to 30 to visit tethered of either sex stituting barrier the develop in estrus dogs ment of normal heterosexual orien when she was in heat (estrus) and tation in adolescence or successful when she was not (anestrus) (Le in anestrus was more functioning as a wife and mother in Boeuf 1967). Visiting fre adulthood. quent during estrus, with a strong preference for visiting males instead Behavioral endocrinology has also of other females. learned much from examination of 15 normal changes in the endocrine Are women sexually most receptive, system and associated changes in attractive, and proceptive during Ov behavior. One example of the kinds the middle of their cycles when of studies that have been undertak ulation is imminent and estrogen en is presented in Figure 1. Here we secretion is high? Interestingly never see that the striking increase in se enough, the question has cretion of testicular androgen which been asked in quite this way, al Tethered ? Tethered in most at about 13 many investigators have begins boys anestrous 9 though years coincides with the sought to determine if "sexual roughly 2. In a of interindividual asso of masturbation Figure study drive" or "libido" is higher at this beginning (average ciations among it was found that fe dogs, and no consistent answer has age 12.0 years), first ejaculation males in estrus ("heat") spend more than time, The situation (13.8 years), onset of nocturnal twice as much time visiting tethered males been forthcoming. as do tethered females we emissions (12-16 years), first date they visiting (Le might be clearer if dealt less Boeuf 1967). This experiment has implica a her own (13.0 years), and first infatuation with woman's analysis of tions for the study of female "proceptivity," These are de feelings and paid more attention to (13.5 years). changes or male-seeking behavior?an area that is or actual sexual as we do in layed absent in hypogonadal beginning to receive attention from behav behavior, 3 boys, who lack normal amounts of ioral endocrinologists. studying other species. Figure

1975 March-April 179

This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions ular hormones are involved in spawning behavior and the preced ing movement upstream, but before these hormones can serve their functions the fish must move from salt to fresh water, and this transi tion depends on the thyroid hor mone. Figure 4, which illustrates the effects of dissolving thyroxin in the ambient water during the winter, shows clearly that the hor mone produced in the nonmigrating fish a preference for fresh water which was lost as soon as treatment , I_l I _L-1I t i I I-1 was discontinued QL (Baggerman 23 2110 17 1613 11 0 7 5 3 1 1962). Reverse cycle day The second dealt with Figure 4. By experimentally altering the 3. Variations in the secretion of es experiment Figure hormonal balance of an organism, behavior trogen at successive in the menstrual effects of adrenal hormones upon points al endocrinologists can temporarily modify cycle appear to be linked to re extinction of a learned avoidance mating the behavior of that organism. In this graph, in sponses human females (Udry and Morris response (De Wied, Bohus, and when thyroxin is dissolved in the ambient Ovulation in the human female is 1968). Greven Rats were first water, the stickleback's preference for salt near 15. As 1968). normally expected day exempli water drops dramatically?a symptom that fied in this behavioral endocri taught to avoid electric shock by experiment, normally precedes spawning behavior and actual recorded behavior over a barrier within 5 sec nology emphasizes crossing the antecedent move upstream. Treatment rather than the of analysis feelings?a pro onds after a buzzer was sounded. was given during winter, when the normal cedure similar to those used in studying After the habit had been learned environment is ocean water. On the seventh other species. day, the administration of was dis perfectly, shock was omitted but thyroxin continued. (Adapted from Baggerman 1962.) the auditory signal was continued; the question was How long would it shows the results of one investiga take for the rats to "unlearn" the tion (Udry and Morris 1968) in acquired response? The experi Examples of hormonally controlled which a small number of married menters found that extinction, or effector structures are numerous in women made daily written reports unlearning, was accelerated by in connection with the reproductive as to their menstrual status and the jection of corticosterone, an adrenal activities of many anurans. At the occurrence or nonoccurrence of in hormone that normally increases onset of the breeding season male tercourse with or without orgasm resistance to emotional stress (Fig. frogs and toads begin to secrete pi during the preceding 24 hours. The 5). tuitary and testicular hormones two shown in curve peaks the upper which induce changes in several are remarkably similar to two mechanisms structures necessary for successful which appear in plots of the fre Mediating mating. In response to hormonal of quency mating responses by We know there are at least two stimulation the laryngeal apparatus some female rhesus at monkeys principal ways in which hormones increases in size, thus facilitating successive stages of their menstrual can influence behavior. One is by utterance of the sex call that at cycles (Michael, Zumpe, Keverne, controlling the original, prenatal tracts females. Brachial muscles and Bonsall 1972). development of the "behavioral hypertrophy to augment the so machinery," to speak, and the strength of the forelimb mating Evidence for covariation as provid other is by modifying temporarily clasp. Cutaneous excrescences ed by pathology and by study of the functioning of that machinery. termed "nuptial pads" appear on normal fluctuation is greatly en the inner surface of the anterior riched and extended by the results The second type of control involves limbs, providing extra traction in of experimental investigations. The what are called "activational" or gripping the female. fruitfulness of experimental meth "concurrent" effects of hormones, ods as applied to problems of co and most of them depend on one or Some male Cervidae, such as the can variation be illustrated by two more of the following types of red deer of Scotland, u?e their ant different quite studies, the first change: (1) alteration of effector lers to compete for social rank, dealing with annual breeding mi mechanisms used in the execution which in turn determines their ac in fish and the second gration with of specific behavior patterns, (2) cess to receptive females. Stags cas in rats. forgetting modification of sensory and percep trated during the mating season tual mechanisms in such a way as lose their antlers prematurely but The three-spined stickleback to influence the reception or inter grow a new pair before the next fall spends the winter in ocean waters pretation of stimuli from the envi rut. However, the new set often is and each spring migrates upriver to ronment, and (3) alteration of cen deformed and unserviceable for in - Prior to the onset of spawn. migra tral neural mechanisms responsible termale conflict. Consequently, al tion, hormone secretion increases in for integration of incoming infor though castrated males can be ren the pituitary, reproductive, and mation or for organization of overt dered normally aggressive by im thyroid glands. Ovarian and testic patterns of response. plants of testosterone, their agonis

180 American Scientist, Volume 63

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'. . krn,_-.;L' ' ...;. ,L,.,.. .F... -,-'ir"i .:,. f.1 .. :. ,_ ,....-,) warning injected,. . I. .: .- . ,1;. : ,-; '. . . i.;L..-.1. . .-. . ..- . ,,. , ,-,; . .. . ; : -.".,;." -;-. ..,..!Q , ,-,;.-..,.--- p -, -S.hp..GP . . -,..-. ;!.:- . ir i".. -,1.;;" . ;.-., . -N ...... -.t - ..--.... --1-,k,.Y,,.k.- . , .. .,..,..,. ; . L, --,,V.- ,J ii.-I, X . MY .- -;. . -- ,-; ;-;,- PIL, ;' '--- .. - -' ' :. .. I ...... , .- ' .L' , . . ..,. .. !, c .L,. ..-A t!'!-,--- _-;'-' .j .e--;. I... . ;UI4A .Ik j4-'i'..'M,*' ,I ?M.., ;0NIT:1 A,-. . ' ..? I . :-- . . ..,- 1: , "Qt'C'L1'J-, .I I .: 1.-1L-Ij V.;- '.. .7m--".' i_-!. - - - with .02 mg of the hormone, and much fast -- -: ..... -,,- -, , , , . . '...... L. .. .,. .:. ! . ,., .., .L. ... ,.. .. -. . .1 ' .. .- . -. L- - ..4 - ,. IT , --;! -, : .i.: ..,I ,N p"1,M -iv. 'In .'. L - - 11.f -%!-TI -Pff . -111-. -ibA;-_ - 'Z:t- - ". ..' :," 'L.-;,'. F,. ,'._L -1 . - ..",.1 I . - - . I ,WO'Itf'. .7, ,,;.- .. 1. , , . . .1,..: .., ...... 7.% . 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- . .'... .. - , i : I - I, . ,. ,, .- .1'...... - .... A- .F;-!,7 ;,.;Im l Y-,; I; Figure 6. Temporary hormonal deprivation has been shown to impair the sensory ca pacities of human females. This graph shows changes in the visual threshold at which a faint light is perceived at different stages of the menstrual cycle. During days 0-5 (menstruation), the ability of normally cycling women to perceive such light is much less than that of men and of women taking pills containing ovarian hormones. (Adapted from Diamond, Diamond, and Mast 1972.)

tic behavior fails to gain for them the elevated social position neces sary for undisputed female associa tion and successful mating (Lin coln, Guinness, and Short 1972).

Hormones can alter not only the ef fector mechanisms but also those involved in reception of stimuli from the environment. In fact, though she probably does not know it, nearly every normal woman who reads this article has experienced changes in sensory capacity caused Mast 1972). Estrogen also increases in the wintering stickleback. Estro by hormones. As mentioned earlier, breast and nipple sensitivity in gen precipitates a preference for before and during menstrual bleed human females, and when ovariec sweet-tasting substances in female ing the body is almost totally de tomized rats are injected with es (but not male) rats, and the unusu prived of ovarian hormones, and trogen, skin areas around the vagi al gustatory yearnings of some one result in most women is a mea nal aperture become more sensitive pregnant women may well have a surable loweringof the ability to to light touch. basis in the shifting hormonal bal make fine discriminations in the ance of gestation. Deficiency in cer senses of hearing, smell, taste, and Instead of changing sensory thresh tain adrenocortical hormones touch. Figure 6 shows the results of a olds, some hormones appear to af evokes a powerful craving for the testof the ability to see a faint light; fect perception and thus to alter or taste of salt in rats and humans, monthly sensory change does not even to reverse preferences for par whereas lack of parathyroid hor occur in men or in women who are ticular forms of stimulation. We mone is associated with a strong taking pills containing ovarian hor have already seen that thyroxin in tendency to ingest substances high mones (Diamond, Diamond, and duces a preference for fresh water in calcium content, such as chalk.

1975March-April 181

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' -!-'iq'.'q ' '6' -N' -';'. -P"':" -4.'... ..'r -:-' 1- . ;..1j.j.',.'L..1:.1',."_ ...... ;!:._q " ..- .-..'."N.': ., - A". '... .." ...... ,-1. 'i ..,..::L ' .1':' ..' ! .--.1L.- ,;..' .% 1 :." "-4iA; ; " - ' 1'...;.IL' .I..'.'.. .'..-- . '. ....::e .-. . q ..q-1..::: - - , .. ... -:-: :..: ..,-.!.S:' '..".;..,gi. L.-'-':-,i; -' ...;...,..v -'-,. ,:-V'- ... S' " ,:.jk --'L '. '4' i ". 'I, 'T. ."..". . .--..?",...--N'R'-R 1.'A ..pA, .*_ I... I.. A ,,24-..'" .!:' J q!'- - -1 I';.-'-...."'-',:-,' .' m 4. ir',..'... %4, . V-..'...S i"--'1-qM-.0 ...to - . mones. Male rats the odor of :' -.. "-# iq". - :- - ... J,_.. --i -'.' 'L:' l..."... , ."-... .. &- ...... " : ,L' I' .1.... ;-!.... m.t i..;..% - -1,1' rk.- i 161AW -1r::4-r - -K .'4V- kl'!;;.'.r ) Vj )!_; -":ti' '-': . -'It-' K4 iff.A 7-11:':Eiz4-'iLijc ,. *..x'-';.. -e .' -, -A ,'-. j ;.-%'."qq'.' ...I.'If. ,.-..' --.i--'q-L"'K".';'%...i :iq... '..I I ,A..':..' _';'.'..L'.,,. . '% prefer t: "L ..; j.' '.L% . , ".,.:..-, .. ...;:' ,-,. .:' L' . . -' ...-'.. ,i,:. .', :...... '... .L..I. .. .q;- ::: .1 ,:",...-.' ::.,t,q;.-i,q.!.- ..,..-.- "W ML Sm... ..- 11ri,w: ...A-q, . .aQ ..I - -'km*-..' -7 -f zq - --.-..,J- .. -j- .. -.'.-_... -'L' ". "... a .t - . ." . .:.v . '-i" 'd.,,. , '..1,-k11 1..'".. "Q.. ..- " i... .. _ r L. i.C: -IL , - '. ..'..':I-q - -'% .,. 1-4: .L 1. ,L.. '. '' a'.,--i' -.:. .. . .;I, - .1.W",A 'O . ."..' 'zw ..'r, I--rr r'R,A .; !.WM1cA q..m. tilz I -'-- i ..'A-'. 41tr-." . .1-j- , ..'4 . . .;..d7-% -L'.N--W._L'L MTF".. ..'L. J ".. .14F 'rx .1. . , . - . estrous to that of females %.... -j.'L.-.... -i .1 ...... 'X ..1 , ':.'---.-: .!...... 14 'Lq'I.':.j.'.' -" - :: ;..' . .. .1: .L ..::.,q !r ' .I!.t:..'' . - ..I.- f".1-1_".. _.'f"Y_ --L:.!;.F' j";' 7 q;'44-.-- .:yj F, -;kp-!g.."'. ! , Ymkn+P. W.. . ":q..t':-* 'L!- ', -V -' 'T.%!'.iq , .I....., .:.... .:.rrI ..ji . females .' 1 '..L' ' .' .- : !-: .. '-"": rL- .-1.' .' .' qq'..-- .;.'. ..'. q -.... ; r .'...... '..- . - ...'.". .i ..-. .-;:_4..e:- m...'.'.- '.4 4 _.z '..; K51103-T.- I, , I'-- ,.-PT',9'. .jgv- .q' ..... A'-k;-i._ '. - .. IL .- --.4w_ .-A4,T.iig-'S -" '" '.L,.--izqw T-- . NP .- -I- -:jv _%,. - -.-''.'k.. '7 .C' -: U. .-4'::!,- 11 ".A ": _' N , - L ; ..1. -.,.'L;'. . .q...-. . ; .L .. "F. , ", q.. - . .- . .1 : . . , ... --.-- .. .:-.4 -..1_1 . . L"1OWN14, ' :_":11-f- _ -'? -' .,.,ff- .' 4P41P. j',, - A 1--W. .14.j.."N. A. - 'q :--. ... .:. .-.' .IM -'t. .r .." m-. .o- ,; - 1,' ..- ..I q..' ii .- . q:: . .: ...ti - - q'.Lj;f.'.J L'&Lf;.. 14"Lti-wei . L.. - C; - . .. . ' LT jF . . r A!'-'..'"- -,L ; . ;'. --.' - - 'L .. . ". . ,'- -.".- -.. -."Yq !' " ...--.. . -,'.. not in and the has . .--'. ... .-' . . --. i'' . .. j - . - - .I . - . I.' -".''.C. ' :-!' 'x. -.'. k- iq.' '-L j .r !.. "' _.', .... -.OXF..- 9?.. '. Zr-'.'-"tl't ---j.' : I.-- 1 . '- -:'-'/Im' .!4'. ."" ..: 7*-! . . p," - heat, preference _:: " .'L.. L '" -' 'L'V -'. - -..-. 7,'-_:- -.-'F .,..: .: ,'. .. . : .. ; .ki,. '.1 q.. . - , .. - .'.' , 1..._ .:.!. . ...r, -.-'-L :6..Ir'lo ,q.w.j X I .-I,,.i.--v' f--.-k.-'k---.T...-il-I 'Nii4'2- -" '-"- -4.4 ' A: m, -, : - 1.".-. .., q.- .- - %, - .. :--I" -", --'. -, -' , 'I .:: . .. ' 1-. - - ,I.2. , : . .- - :z ; 4: %L.7 .-..-- 9., q:prg't v -,' _Y' .-.: ..'. A.,,A ...... shown to on testis ..' .' ...... i .. '1.k -- -!-.'" - .q . . . -" ;1- -' ';- W4;'41 - , .;, .. g. ...-..'..'I. . I -_---' - 'L e' ;L;' ..'-!-Lf. -1- I.e.q..., ;;,% 4 1.11'.4.'.'.-.. .' .1 -_-4"'f: been depend ;1- ... .-. r .., ..:. ... .' ' . .-I ...... t:'- ..,;..% :., m . - i ; .. ,. 1.;::. , :. . , - ..'A".. ''C,.f.'.% rl t. 4;,j, ' ' .1 ...'; &&lE,.t;tL I .._.: . M . . ., ""Wi.. : - . ...' ..- ...... '.1! :.;: -- m- ':r'' ... .X- .. . .1. ,:;.. . -fF!.. .:'L.". -: - .- .- .; , :.-- .. .. '-j - c -I ". - -- q.-:!_ f;Ur -h.;F.- .;'!' .?w F_.... .- .. ..p ..")4:1-. ,- - A : . .:'. .- ;'.I- -S- - ._ -'r- , . ';-.. ..., - . . , ,'? .-"' '... C. . ...1, _.- ". ".. .. , ...... q - - - - .;-.-.;-.Z.'!!Lj ,. - -4, -i'<' -',,, . - . .11,.11I-.1 mL-. ' ..;... .'. I ' - -F. N-i:A.A .:: f& ".":-1 : - P- -..-*' k 4;q.1.....- P'f. 4- --1,"t.,IA .' r! 'm. '- ';F --, ..';;j;j; ' fL hormone in the for it can be -':..-i...: : :. -.:" .- %;., , -. -.,'-, .. ,'- '. ..'t.. !L; ," : -....: .1, .-'." . ,.1. : :'-4.J,- . '- - . .- :. , . ., .&': . ..'A' I"d -f" - m-W- ,M. m 1-..'N ' -' -5'-L -I. . :.--A, ..4, A S., r ' Lip' 'JF..- - ". ;.% ".1.1 .'. -TT'. . IIt.'; 1, 'e4Y-'k- Xr , .'. I .i,..;;: .' male, j' --i7..r.-. : "L r -'__' -'- I,,- - - -."' .- L-- .. !-' , . . . .-n. -'.:!..- 11.il'K - ...... --Oq '"I''d..-.7 .1.9f"' " . . ,' I -it" 1_. ;j- -MlmV . --... z ;iN ,--) - ....1. _&'... , -t-:. _, 1 ..m -'L.""L;.,; : c -, _" :'i'.J - -A' -O 4, L'..Pf I ., .. ' -7..' ' ....:...-Z.r..-I., ,-k...': 4-.., ::. - -: .. ..L.'I.. 'y I Ik.".. ' - .'j ,'It','"" -''.._''.' "I"- .q. ,-.-' . . . -I'...... -. ... t.. 1.' L- L,.;? q'F1731-'.1.11 1.4.11- _;'-' rg .'. '.Z r jif.;%L. . , 1,- -., - ,I " ,q-. and then ! , -.; , ...... z1:. ... -.. '!'r .." ,L '-"-- _ i :--':t.'. .1. 71'i - . - .e.L.f".. .. : '..,...... ,11 , .- . ,- , . '-.. 1;;.,;r", -'. ,'.q : ,...'p'M I -' -"'.v ".T4 j'.-. .-';- ;L' . - e., .!_;, '4'..A -.!-'."'T. -!;-T .L'_t".'.' t'i'141-11-Kt "'A ..-.-._ , " , f_.. 1. eliminated by castration 1'.: -'.-6.1 I-' - -:.- , . .' LV ',. F, ;_. -.' - i.' ... .'.k ' -,." ..1".4. . ,t::! t. -. .'.f,. mq . -,,, '.. :.".q . ov,Wei ' ZL'; W..N. 11. I- ";Q.FjLS' . P-..--. f-"- pr.--.'vv -" .i-* . --i va.1 Ap --'* .1-v - - f- -1. ','1041'.-, ..-.: - -- .. ..L. . I1. -, -I -r . -'.--; --.; . '.; . .,: .: %'-- ...k::. ,L '-'.- - ...... -: 1...j i..9;' 4..-.15"', k .,j - 'A I---, o'. ' L.L-'-'...'::'i.'; .W.,'T'.'. M'-.; 11'.'? .I .;'W li. -1;710-a'. .b .. . ! . , .'. I lr-: 4p. -- ;Z..-' .-, .. ,,I '% 1..n'.. . ..1.:.2.. , ...... 1 ' '.".... - I- I t...;,;, q . ' ... .: .r"- ...- k -141:' , J-r-. '. , -,F-. w., Lx. Lf ..t_: , , .-'qp. tallf.-k- -.?'0M '. .1, I ; restored testosterone 'i..' . ,.4. .. Iq.A ...... - ..- ...... '.d - L'-..' -,-V!. -A q.;...; ;_ -'%- .1-'.:4 NP.-I ...64Z., 7. -1 a ;;v"I":- TVI.; _ . 5 z'ift . ).5-.11 - . .' '!%' .! .. cU.. ,-.:L.JEj: X "q--11:"r ..' I. .4 . 1..:g,:. , by injections. . : , , ...... ,,, "".iz % - .-.-c. ... - ...-_'w: 1'. j- - -." - q ..-"-- - .' !, 41I.j.':'r, ' .," . . . . I.; .l", v"..?-.1..J.. '. ..%. Ni ..'. '.. .. q.;.- .....-' ... - p....' - - ..r. "., , ...... -.Lj ...... i:- ,: L ,'t! . .;:"_;. .- e'--. ";. : ....-. .'- '.._.-';;rw" I .. J'--A..'f , "- s. 0fi ".,k- 'r -I.-ILA r_-....7 .KC -17., .7 ' 'L OC' . y;. ff ,- _ C_-.-- - are .. .; . .. ..C'.,. :. . . . _I , -L .!.Z '7 , ..:, . q -_;- 'O. q, 'q! %I...- - f- 117T1. i .j -L-1 1.- A'k, 0 -'L. 0.'-'*'.L.&.jv "10 ..'CL L i - i; %.J.Lj--- , ;q.-J... -!M '4p'nP1:q..j -q.&P'._P. Q !R'1- - Even after they castrated, male .. . . L,. ...I. . . .qq%. . ix.:.',..q ...... !- . . . ..i....- al' . :r: -.. .. f : . . . I : .:"q ..,q -;'! . .,;-- --.1: .-::% --' - ;-';'V' -,I .';..11-.1; - . 1tyq7.', .Z"-': i -';i4f'r4".-.'.jr ,, -' - .-.. - -.'-.-.'w ---4k-'-F" q; O! . q--.- '. ."f'..J. j'r2 .--,';Al". . p! . . . ,:.LS ', ;_- _- i. :% - . 4. .:1, :- ..1 -. , ' ..'Lq..i". -. . .L"- ..'. ; .'_ .'- : '.. -. ..." ., ... . -,...... ,S.-, .. .'.'. ..- ,,:, .'J.,'rY.; j . . . . t!1.I. '. -- .'. 1.5- ...- -4, - "I - - . -.L 0k . :.--x.'i ..1114m1 t. %--':A jo 1. : ... . . qf1. ... -.' ::"".'--)I,,.. :1I -. :...... :, .'----'!%!.-J.1, ....' ..- ..- ..:.- . , I .- . . :I -% -' !' .....;,. I - '.L..'...... -. -.. ','_. 0_ ...q : L. 1. -.1 . ... q.. b .-'!% q . ... .'-. ..;fT.', I' ..- ,.... .". - ' ...... --j-v?-..'.'-'O!'.-.-.-: .:,.c.... q..". a for ...... : . . m -.1-.m.,q.". I- - I . .1 . . 1L.-- q . 1,q), --' .. '.: ...... '. qi .N. i...... 11 F;v r. I., , * .. " .. q ..rq ! !"- .'. - -4,- C. . -'..o .".:.;.-Lv-. ...;:.. '!!" L%' W1%.. .-'1 I~ .. ;.. ...f-q.,... Alk"'.0(_.W. f - 11.1. dogs display slight preference . . . . _. . .'-- v - .'.;i__; , %....- .; . vaginal odors of estrous versus an estrous bitches, but the preference is much stronger in normal males and is increased in castrates if they are injected with male hormone.

Female dogs can discriminate be tween male and female urine and can even bUstinguish urine of a fe male in heat from that of one not in heat. Spayed females and normal males alike show a preference for the odor of estrous female urine, but when the spayed bitch is brought into heat by estrogen injec tions her perception is altered and she now prefers urine from males to female urine of whether regardless Figure 7. Among the known developmental with testosterone, the male hormone, were the urinating female is in estrus or effects of hormones is the inducement of observed to exhibit a greater frequency of anestrus. sexually dimorphic behavior. In one experi rough-and-tumble play than did normal fe-* ment, 12- to 15-month-old female rhesus males of the same age. (Adapted from Goy monkeys that had been treated prenatally and Phoenix 1971.) Despite a plethora of evidence proving that hormones can affect behavior by modifying effector and receptor functions, the mediating mechanisms most often studied and emphasized have been those attrib uted to special areas in the brain. It come sexually receptive to males The last decade has seen an up is clear that endocrine secretions after estrogen implants in the hy surge of experiments on develop can modify brain function in a vari pothalamus (but not in the cerebel mental effects of hormones in a ety of ways. This has been estab lum); and male rats which cease number of animal species. In lished in part by implanting elec mating after castration will resume species with a very short gestation, trodes in different brain regions and copulating and ejaculating fol such as mice, rats, and hamsters, monitoring neural activity while lowing placement of testosterone in many of the changes described can the animal's endocrine status is al the anterior and be induced by direct administration tered experimentally. This tech preoptic region. of androgen to newborn females; nique has revealed that stimulation but the most extreme effects are of the vagina and cervix, such as In distinction to concurrent effects produced by prenatal or prenatal would occur during intercourse, al of hormones on behavior there is a plus neonatal treatment. The gen ters neural activity in the limbic re second category, known as "organi eral trend of results can be summa gion of the female rabbit's brain, zational" or "developmental" ef rized as follows. When pregnant but only when the brain is simulta fects, which are distinguished by rats, hamsters, guinea pigs, dogs, or neously stimulated by estrogen. In three characteristics: (1) they are monkeys are injected with andro contrast, a different ovarian hor permanent and irreversible, (2) gen, their female offspring are mod mone, progesterone, tends to de they normally occur exclusively ified in several ways. They are born press the responsiveness of certain during special stages of ontogeny with masculinized external sex or neurons in the brain of the female such as the prenatal or neonatal gans but with internal uteri, tubes, rat. periods, and (3) they are not mani and ovaries. In addition, their ten fest at the time of their occurrence dency to exhibit female sexual Another way of investigating effects but only become apparent much traits is markedly diminished?i.e. of hormones on the brain has been later in life. Developmental effects they are partially defeminized? to implant minute quantities of are known to influence bodily while at the same time their incli crystalline material directly into se structures, brain anatomy and nation to behave like males is in lected nuclei. This has shown that physiology, and sexually dimorphic creased, indicating some degree of ovariectomized rats and cats be behavior. masculinization, which is an entire

182 American Scientist, Volume 63

This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions males when it comes to influencing selectedfor high HH __| secretory activity of the pituitary un randombred gland. It is very important to H? that of sexual selectedfor low derstand plasticity IJjjjjj^J mechanisms in the rat brain is re stricted to the first few days of postnatal life. The reversals just described cannot be induced if cas tration of males is delayed until the seventh day of life or if females are given testosterone at 16 rather than 4 days of age.

Contingency in covariation

Functional correlations between hormonal and behavioral variables depend upon a variety of factors, the most important of which are the genetic composition of the orga nism, its current state, and its per sonal history.

Although the same hormone, tes tosterone, affects vocal behavior in Intact 6* Capon Capons given androgen the male bullfrog, alligator, robin, and moose, the sounds produced by 8. differences in the same amount of testosterone, those with Figure Although qualitative these animals are vastly different. a from a selected for the effect of particular hormone vary genotype high mating frequen are of more twice as often as These qualitative differences species to species, genetic differences within cy copulated than a for low course caused differ a species can influence quantitative effects those with hereditary tendency by underlying of that hormone. When capons from 3 ge mating frequency. (Adapted from McCol ences in species genotypes. Genetic netically distinct lines were injected with lom, Siegel, and Van Krey 1971.) differences between different strains within the same species can influence quantitative aspects of hormonal effects, as shown in Fig ure 8, which illustrates the effects of castration and subsequent tes as as of tosterone on ly different process from defemini well the genital anatomy injections mating by dis zation (Beach 1971). female animals. The electron mi roosters from three genetically croscope reveals very fine differ tinct strains (McColIom, Siegel, Strain differ Evidence of such changes can ap ences in the connections between and Van Krey 1971). one ences are obvious in normal pear in many types of behavior. For certain types of nerve cells in which area brain in newborn birds are almost obscured cas example, as illustrated in Figure 7, small of the by at least some tration and when the play of immature female mon males and females of (caponization), that the hormone is in keys which have been prenatally species. It has been shown given equal can or amounts to all those with exposed to male hormone is more these differences be reduced castrates, like that of littlemales than is the eliminated by very early hormone a genotype selected for high mating more than play of normal females (Goy and treatment (Raisman and Field frequencies copulate as as with a he Phoenix 1971). Adult female dogs 1973). For example, if a female rat twice often others which have been stimulated by an is injected with male hormone reditary tendency for low mating There is clinical evi drogen in utero and early in infancy within 4 days after birth, her brain frequencies. so that it dence which that human display weak and infrequent femi is permanently changed suggests a inherit individual differences nine mating responses and are not assumes the appearance of male beings to attractive to males even after ad brain; and during adulthood in its in their sensitivity psychological ex effects of hormones ministration of ample amounts of control of pituitary activity it specific (Turner and estrogen. On the other hand, such a hibits functional characteristics Bagnara 1971). bitch is much more likely than nor typical of males rather than fe to mal females to exhibit masculine males. Conversely, when male rats There is ample evidence prove that the individual's copulatory reactions toward anoth are castrated on the day of birth physiological a time affects behav er female in heat and to lift one they are deprived of the testoster state at given ioral to hormones. For hind leg while urinating. one which normally influences de responses the of testosterone velopment and their brains are mi example, effects treatment on the behavior of cas It is not entirely clear why prenatal croscopically like those of females. trated to the exposure to androgen should mas Furthermore in adulthood these stags vary according culinize or defeminize the behavior brains function like those of fe time of year the hormone is admin

1975March-April 183

This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions istered, because the animal's entire of masculine sexual behavior in fe own and all other species, insisting of hormones in physiological balance undergoes male rats reveal that females whose that possible effects never accu seasonal changes. When androgen location in the mother's uterus dur Homo sapiens can be was because inevi pellets are implanted in December, ing pregnancy between two rately assessed they as are masked or mating responses appear in a few male fetuses are five times likely tably overridden by weeks; but identical treatment in to show masculine mating re effects of psychological variables as are such as and April does not elicit rutting until sponses in adulthood females "expectancy" "sugges five months later, in September, whose intrauterine position was be tion." This "human chauvinist" which is the normal mating season tween two other females (Clemens position is doubly disproven by the have seen for this species. Annual rhythms of 1974). One possible explanation is evidence. We already of hormones on responsiveness to hormones are that testosterone secreted by the that many effects seen in females as well. Thus the sex glands of a nearby male fetus human behavior are well estab ram-seeking behavior of spayed can diffuse over short distances and lished, and it is well known that can ewes is easily evoked by progester partially masculinize a female lit personal experience modify be in one and estrogen treatment in the termate. If this were true, it would havioral response to hormones fall, but females are quite refracto be most interesting to study psy many nonhuman species. ry to the same hormones during chosexual characteristics of human of the early spring. females with twin brothers. Rather dramatic examples effects of social conditioning upon have been A person's temporary psychological Other experiments have shown that reproductive behavior in studies "expectancies" may affect his re when a pregnant female is repeat discovered by ethologists sponses to hormones, as has been edly exposed to emotional stress, of what is termed "sexual imprint shown by an experiment on the the consequences may include per ing" (Klinghammer 1967). If eggs of are to emotional effects of injecting epi manent modification of the behav certain species of birds given a the nephrine or adrenalin (Schachter ioral responses of her offspring to adults of different species, them and and Singer 1962). A number of con hormonal stimulation (Ward 1972). foster parents incubate rear ditions and subject groups were in Male rats born to mothers systema the young. When males which cluded in the total design, but only tically stressed through the last have matured under such condi a few are relevant to the present third of gestation (by periodic ex tions become reproductively active, discussion. At the time of hormone posure to bright light) were tested they engage in courtship and copu administration some subjects were in adulthood for the display of bi latory behavior as do normal males a told what physical symptoms would sexual mating patterns. When they but show strong preference for the foster be produced, whereas other groups were injected with testosterone and mating with females of own were given no information or were placed with receptive females, the species instead of those of their deliberately misled to expect false experimental animals proved to kind. Apparently social learning not affect symptoms. While the hormone was have been somewhat "demasculin during development does to to testis taking effect, half the people in ized"?their display of mounting the capacity respond each group were exposed to a con and ejaculatory responses was infe hormone, but it does determine the trived social situation previously rior to that of normal males. When stimulus ("sex object") that will shown to arouse euphoria, and the they were injected with ovarian evoke courtship and mating. Here others to a different one intended hormones and tested with stud we can detect at least some formal to evoke anger. Susceptibility to males, the experimental subjects similarities to certain psychoana manipulation of the emotions was showed evidence of feminization? lytic theories concerning early life increased by epinephrine in those that is, they executed more female experience in human patients and subjects lacking foreknowledge of mating responses than did control homosexual object choice in adult its physiological effects; but those males given the same estrogen and hood. who knew in advance what physical progesterone treatment. One possi symptoms to expect were protected ble explanation is that stress during Interactions and the intended emotional ef pregnancy elicits excessive secre against interdependencies fects of social suggestion. tion of mildly androgenic adrenal hormones, in both mother and off A condensed review such as this The third category of contingencies spring, which compete with the might easily give the impression involves those which are rooted in more potent testis hormone, pre that behavioral endocrinology deals the subject's personal history, in venting it from* completely mascu primarily with one-to-one relation cluding the effects of learning and linizing and defeminizing the ships between hormones and be experience as well as those of physi brains ofmale fetuses. havior; but in fact the fourth and re ological stimulation. Examples of possibly the most important developmental effects presented An entirely different dimension of search area deals specifically with earlier dealt with human pathology personal history that affects behav interactions and interdependencies or with experimentally induced ioral responses to hormones is that among the endocrine system, the changes in animals, but it is possi encompassed by such terms as indi individual's environment, and his ble that normally occurring hor vidual experience, learning, and so behavior. The practical complexity monal variations in the prenatal cial conditioning. The importance and theoretical reach of the prob environment may have lifelong ef of this sort of contingency in lems constituting this category can fects upon the behavioral response human behavior is so marked that be illustrated by examples repre to hormones. For example, studies some authorities dichotomize our senting 3 subdivisions of the area.

184 American Scientist, Volume 63

This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions 55 secrete testosterone and produce spermatozoa.

In several mammalian species in cluding our own the activity of 50 some endocrine glands can be in hibited by environmental stimuli. A woman's menstrual cycles may be abolished for months as a result of emotional shock, such as oc curred repeatedly during bombing 45 raids inWorld War II. Secretion of testosterone can be temporarily suppressed in healthy young men exposed to psychological stress such as that induced by rigid require 40 ments of military officers' training programs. In a recent study of changes in testosterone secretion by two male monkeys in different so cial conditions (Rose, Gordon, and Bernstein 1972), both males were 35 placed with groups of females after Control 9 exposed from infancy to: two weeks of social isolation, and & behind wire & bedding castrated & hormone secretion increased promptly. Subsequent transfer to 9. Environmental stimuli, normal males have been found to reach Figure including pu all-male groups, in which each ex those provided by other individuals of the berty at a much earlier age than females not can male was and same species, affect the functioning of exposed to males or those exposed to cas perimental severely, As in the endocrine system. shown this trated males. (Adapted from Vandenberg successfully attacked, resulted in graph, female mice exposed to odors from 1969.) lowering of testosterone output, which remained depressed for ten weeks until a second exposure to fe males brought the level back up again.

The secretion and release of hor a male viewed through a glass plate The maternal responses of mam mones by various glands is far from is enough to accelerate growth of mals depend in part upon prolac an automatic or self-regulating pro eggs in the ovary and secretion of tin, which is produced by the ante cess. It is subject to a galaxy of en estrogenic hormone. In some other rior pituitary gland. As long as the dogenous and exogenous controls, types of birds exposure of the fe young depend on mother's milk, among which are various forms of male to the sound of the male's continued secretion of prolactin is stimulation received from the ex song provides an important audito facilitated by stimuli the female re ternal world. For example, certain ry stimulus for ovarian responses. ceives while the young suck on her hormonal rhythms are effectively This general type of environmental nipples. If the young die or are re synchronized with specific changes control appears to have evolved in moved, prolactin secretion soon the in physical environment. Thus dependently in several vertebrate stops and maternal behavior disap the springtime increase in length of lines, for ovarian responses to social pears. If, however, a female's own day triggers secretion of pituitary stimuli are known to occur regular offspring are periodically replaced and other hormones essential to ly in several species of fishes and by younger foster litters, the resul successful reproduction in some reptiles as well as in birds and tant stimulation of nursing can ex species with a spring breeding sea mammals. tend the period of lactation and son. In other instances effective en maternal care to many times its vironmental stimuli may be provid Males as well as females are open normal duration. ed by other individuals of the same to control of hormonal secretion by species. As illustrated in Figure 9, social stimulation. Rhesus monkeys In addition to the effects of envi immature female mice exposed to living in nature breed for five ronmental stimuli, the individual's odors from normal (but not from months of the year and are repro own behavior produces various castrated) males reach puberty at ductively nonfunctional during the kinds of feedback effects leading to an earlier age than control females remaining seven. Females captured changes in his endocrine system. which have not been stimulated by in the nonbreeding period were The simplest or "short-loop" type male odor (Vandenberg 1969). brought into estrus by injections of of feedback occurs when execution estrogen and housed in cages near of a hormonally conditioned behav Facilitation of ovarian activity by captive males which could smell, ior pattern gives rise to stimuli common social factors is in birds as see, and hear them. Nonbreeding which more or less immediately de well as mammals. In the case of the males responded to stimuli from crease or increase secretion of the female ring dove, the mere sight of the treated females by beginning to very hormones that were responsi

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This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions ble for occurrence of the behavior in Lehrman and his colleagues at the the first place. For example, endo Animal Behavior Institute, Rutgers Anticipated crine changes during the latter part University. A highly condensed de developments of pregnancy cause female rats to scription of the essential steps in a engage in frequent self-licking be the concatenated sequence follows. An emerging discipline and grow havior which is concentrated in the ing child have several characteris a sex to nipple area. Licking stimulates When a female is placed with tics in common. Growth tends nerve endings in the skin, which ually active male the visual, audi occur at an uneven rate, and phases leads to an increase in secretion of tory, and tactile stimuli provided of relatively slow change alternate the hormones that are responsible by his presence and social responses with shorter periods in which the for mammary development and for to her induce the growth of eggs in child seems actually to "shoot up" or to postparturitional care of the young. the ovary and secretion of estrogen. physically mature very sud denly psychologically. Behavioral The simple execution of a specific Estrogen inclines the female to in endocrinology is currently undergo behavior pattern can evoke proprio creased cooperation in courtship ing just such a growth spurt after ceptive stimuli which in turn elicit and copulation and also facilitates several decades of relative quies Nest cence. important hormonal responses. Re the initiation of nestbuilding. lease of eggs from the ovary into ing behavior is augmented by pro the oviduct does not occur simply gesterone, and the activity of nest Growth is not balanced or synchro because the ova are mature and building gives rise to stimuli caus nized in such a manner that all or ready for fertilization. In several ing the pituitary to release the lu gans or systems develop at the species of birds and fish Ovulation teinizing hormone, which in turn same rate and reach comparable is delayed or completely inhibited induces Ovulation and egg-laying. levels at the same time. A definite until the female can go through the unevenness in the advance of be motions of constructing a nest. Pro As one consequence of Ovulation havioral endocrinology is reflected more viding her with a nest already built the ovary secretes progester in the excessively large proportion con and ready for use does not provoke one, which intensifies the female's of research to date which has Ovulation, because the essential tendency to incubate the clutch by centrated upon sexual behavior and stimulation derives directly from sitting on the nest. As she incu gonadal hormones. This state of the activity of nestbuilding. She bates, the female receives tactile imbalance undoubtedly will be cor more must perform the act of building and thermal stimuli from the eggs, rected gradually as effort is before the pituitary hormone that and these provoke release of prolac channeled into study of hormonal in non triggers Ovulation is released into tin by the pituitary. Prolactin contributions to learning, to the circulation to act on the ovary. turn stimulates the "crop sac" in sexual social behavior, and to a va the wall of the esophagous to pro riety of homeostatic behaviors such In other animals?notably such duce a substance known as "pi as feeding, sleeping, and adjusting mammals as the rabbit, cat, and geon's milk," which will be fed to to acute and chronic forms of several rodents?the release of pitu the squabs as soon as they hatch. stress. itary hormones responsible for Ovu lation is set off by stimulation of When, under the influence of prolac Periods of rapid growth and matu the vagina and cervix, which nor tin, the female feeds and incubates ration sometimes provide the clear mally occurs during the act of co her newly hatched young, the per est insight into potential for future pulation. Secretion of various hor formance of her maternal duties achievement. It appears reasonably mones sometimes is increased in gives rise to stimuli that feed back certain that behavioral endocrinol males by particular forms of sexual to the central nervous system and ogy will continue to expand, both behavior. Within five minutes after contribute to continuation of pro as an applied and as an experimen a bull mates with a cow there is a lactin secretion. tal specialty. We have only begun fivefold increase in blood levels of to comprehend the many ways in are the pituitary hormone which stimu As the squabs grow stronger and which hormones affect and lates testosterone secretion; and the cease needing or seeking regular affected by human emotions and be concentration of testosterone itself feeding by the female, she is de havior. is elevated shortly after coitus in prived of stimuli which the young to science male rabbits and possibly in human formerly provided. Eventually, loss With respect biological males. of stimulation from the young leads as a whole, and particularly to tra to shutting off of prolactin by the ditional endocrinology, it is abun of One final class of interactional phe pituitary, and that gland now shifts dantly clear that explication nomena included in this general to secretion of the "follicle stimu functional relationships involving problem area involves a form of lating hormone," which causes re behavior, the endocrine system, the nervous environ "chaining" or concatenated inter sumption of the growth of eggs in system, and the our un dependency best elucidated by spe the ovary and consequent secretion ment will greatly increase role of hormones cific example. The most thoroughly of estrogen. At this point the fe derstanding of the analyzed case is that of reproduc male is ready to commence another as agents in evolutionary survival tion in the female ring dove, Strep full cycle of courting, mating, nest and change and in the individual's topelia risoria, which was exten building, egg-laying, incubation, adaptation to the demands of daily sively investigated by the late D.S. and rearing of young. existence.

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This content downloaded from 147.26.169.56 on Tue, 21 May 2013 16:09:08 PM All use subject to JSTOR Terms and Conditions arousal levels during the menstrual cycle. Michael, R. P., D. Zumpe, E. B. Keverne, References J. New. Ment. Dis. 155:170-76. and R. W. Bonsall. 1972. Neuroendocrine factors in the control of behavior. Goy, R. W., and C. H. Phoenix. 1971. The primate In?ec. Horm. Res. 28:665-706. effects of testosterone adminis Progr. August, C. P., M. M. Grumbach, and S. L. Propionate tered before birth on the of Money, J., and A. A. Ehrhardt. 1968. Pre Kaplan. 1972. Hormonal changes in pu development behavior in female rhesus mon natal hormone exposure: Possible effects berty: III. Correlation of plasma testoster genetic In Steroid Hormones and on behavior in man. In and one, LH, FSH, testicular size, and bone keys. Brain Endocrinology ed. C. H. and R. A. Human Behaviour, ed. R. P. Michael. Lon age with male pubertal development. J. Function, Sawyer don: Oxford Press, 32-48. Clin. Endocrinol. Metab. 34:319-26. Gorski. Berkeley and Los Angeles: Univer University pp. sity of California Press, pp. 193-202. Raisman, G., and P. M. Field. 1973. Sexual Baggerman, B. 1962. Some endocrine as dimorphism in the neuropil of the preop pects of fish migration. Gen. Comp. En Hamburg, D. A., R. H. Moos, and I. D. tic area of the rat and its dependence on docrinol. Suppl. 1:188-205. Yalom. 1968. Studies of distress in the neonatal androgen. Brain Research 54:1 F. A. 1971. Hormonal factors con menstrual cycle and the Postpartum peri Beach, 29. od. In and Human Behav trolling the differentiation, development Endocrinology ior, ed. R. P. Michael. London: Oxford Uni Ramsey, G. V. 1943, The sexual and display of copulatory behavior in the develop - Press, pp. 94-116. ment of boys. Amer. J. Psychol 56:217-34. ramstergig and related species. In Biopsy versity ed. L. R. Aron Rose, R. M., T. P. Gordon, and I. S. Bern chology of Development, Kephart, W. M. 1973. Evaluation of roman son and E. Tobach. New York: Academic stein. 1972. Plasma testosterone levels in tic love. Med. Aspects Hum. Sexuality the male Press, pp. 249-96. 7:92-108. rhesus: Influences of sexual and social stimuli. Science 178:643-45. F. A. 1974. Behavioral Beach, endocrinology E. 1967. Factors Klinghammer, influencing and S. E. 1962. and the study of reproduction. Biol. Rep Schachter, S., Singer. Cog choice of mate in altricial birds. In Early rod. 12:2-18. nitive', social and physiological determi Behavior, ed. H. W. Stevenson, E.H.Hess, nants of emotional state. Psychol. Rev. Clemens, L. G. 1974. Neurohormonal control and H. L. New York: Rheingold. Wiley, 69:379-99. of male In sexual behavior. Reproductive pp. 5-42. C. J. T. Behavior, ed. W. Montagna and W. A. Sa Turner, D., and Bagnara. 1971. LeBoeuf, B. J. 1967. Interindividual associa dler. New York: Plenum Press, pp. 333 General Endocrinology. Philadelphia: tions in Behaviour 29:268-95. 65. dogs. Saunders.

De Wied, D., B. Bohus, and H. M. Greven. Lincoln, G. A., F. Guinness, and R. V. Udry, J. R., and N. M. Morris. 1968. Distri Short. 1972. The in bution of coitus in the menstrual 1968. Influence of pituitary and and reno way which testoster cycle. cortical hormones on conditioned avoid one controls the social and sexual behav Nature 220:593-96. ior of the red deer ance behavior in rats. In Endocrinology (Cervus elaphus). Vandenberg, J. G. 1969. Male odor acceler and Human Behavior, ed. R. P. Michael. Horm. Behav. 3:1-9. ates sexual maturation in female mice. London: Oxford Press, pp. Endocrinol. 84:658-60. University McCollom, R. E., P. B. Siegel, and H. P. 188-99. Van Krey. 1971. Responses to androgen in Ward, I. L. 1972. Prenatal stress feminizes Diamond, M., A. L. Diamond, and M. lines of chickens selected for mating be and demasculinizes the behavior of males. Mast. 1972. Visual sensitivity and sexual havior. Horm. Behav. 2:31-42. Science 175:82-84.

"This one writes some fine lyrics, and the other one has done some beautiful music, but they just don't seem to hit it off as collaborators."

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