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Habituation Of Aggression In The Three-Spined Stickleback (Gasterosteus Aculeatus L.)

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Habituation Of Aggression In The Three-Spined Stickleback (Gasterosteus Aculeatus L.)

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Content This dissertation has been microfilmed exactly as received 6 7-419 PEEKE, Harman Van Slyck, 1937- HABITUATION OF AGGRESSION IN THE THREE-SPINED STICKLEBACK (GASTEROSTEUS ACULEATUS L.). University of Southern California, Ph.D., 1966 Psychology, experimental University Microfilms, Inc., Ann Arbor, Michigan HABITUATION OP AGGRESSION IN THE THREE-SPIKED STICKLEBACK (GASTEROSTEUS A C TJLEA TU S L .) by Harman Van S lyck Peek® A D is s e r ta tio n P re se n te d to th e FACULTY C O P THE G R A D U A TE SCHOOL UNIVERSITY OP SOUTHERN CALIFORNIA In P a r t i a l F u lfillm e n t o f th e R equirem ents f o r th e Degree D O C TO R OF PHILOSOPHY ( Psychology) A ugust 1966 UNIVERSITY OF SO UTHERN CALIFORNIA T H E G R A D U A T E S C H O O L U N IV E R S IT Y PA R K L O S A N G E L E S , C A L IF O R N IA 9 0 0 0 7 This dissertation, written by Harman. _ V an,. S l z c k .. P e e k e ................ under the direction of hT3...Dissertation Com mittee, and approved by all its members, has been presented to and accepted by the Graduate School, in partial fulfillment of requirements for the degree of D O C T O R OF P H IL O S O P H Y e ............ Dean />„, A ugust 1966 'hairman ACKNOWLEDGMENTS I would lik e to ex p ress my a p p re c ia tio n to my d is s e r ta tio n com m ittee, P ro fe sso r W yers, P ro fe sso r S lu c k ls and P ro fe sso r Savage, f o r t h e i r aid d u rin g the ten u re of my candidacy fo r t h i s degree* p a r tic u la r g ra titu d e i s expressed to P ro fe sso r E v e re tt J* wyers fo r h is encouragement and su p p o rt throughout th e p erio d of my graduate study* H is ever p re se n t a v a ila b ility f o r d isc u ssio n and encouragement of in d iv id u a l i n i t i a t i v e in th e fo rm atio n and im ple m entation of re s e a rc h p ro je c ts was p a r tic u la r ly h e lp fu l* A s p e c ia l debt i s owed my w ife f o r p a tie n c e In face o f my discouragem ents and disappointm ents which she fre q u e n tly had to shoulder as a d d itio n a l w eight during h e r own graduate tra in in g * W ithout h e r su p p o rt t h i s d is s e r ta tio n might nevor have been done* F in a lly , I would lik e to thank my m other, Mrs* H elen Peeke, and my g ra n d fa th e r, Mr* Fred T u rn er, who never h e s ita te d to o f f e r t h e i r encouragement and f in a n c ia l a s s is ta n c e throughout my grad u ate tra in in g * 11 TABLE OF CONTENTS Pag© A C K N O W LED G M EN TS............................................................................. i l LIST OF TABLES............................................................................ v LIST OF FIGURES........................................................................... v i C hapter I . INTRODUCTION . . * I II* BACKGROUND.............................. k The R eproductive B ehavior of th e Three-* Splned S tic k le b a c k • •••••« *• • I 4 . A ggressive B ehavior D uring th e Repro d u c tiv e C ycle * * * * * 9 I I I . PROBLEM........................................................................... 1$ H a b itu a tio n , F a tig u e and Sensory A dapta t i o n ........................................................................ 15 H a b itu a tio n : M ethodological and Con c e p tu a l C o n sid e ra tio n s • * * • • * • • 17 S tim ulus I n te n s ity and H a b itu a tio n R a t e ......................«• «• • * * • •• •• 17 Long Term Temporal P e rs is te n c e • • • * • 18 Components of a B ehavior C lass • • # • • 16 D isplacem ent A c tiv itie s * • • • • • • • • 20 E th o lo g ic a l T heory: D rive I n te r a c tio n , D isplacem ent and th e D is ln h ib itlo n H ypothesis * • ................... •• «• • •• 23 An E x te n sio n of th e D is ln h ib itlo n Hypoths s i s • • • • • • • * * • • • * * 27 H a b itu a tio n as a Source o f I n te r n a l I n h ib itio n • • • • • • • • • • • • # • 3 H > Summary and P re d ic tio n s * • •• • • • • * 3o IV* METHOD........................................................................... kO P r e te s tin g * ........................... * .......................... I 4 .3 The Sex T e s t .......................* * * lj-3 l i i I Chapter page E xperim ental T reatm ents. lj-5 A ggression E l i c i t i n g S tim u la tio n Groups • • • • • • • • • • • • • • • « L6 N on-S tim ulation Group. .............................. $0 P o st-T e sts . . . . . . . . ........................... 50 S u b je c t Assignment . . • • * . • • • • • 52 Dependent V a ria b le s . • 52 D rive In d ic a to r V a ria b le s. • • • • • • • 53 R ecording and Measurement of Dependent V a ria b le s ..................... 55 V. RESULTS.......................................................................... 57 Group E f f e c ts . 58 In d iv id u a l S u b je c ts . • • • • • • • • • • 7& V I. DISCUSSION 119 H a b itu atio n o f F ixed A ction P a tte rn s • . 120 The D is ln h ib itlo n H ypothesis • • • • • • 128 The E x ten sio n o f th e D is ln h ib itlo n Hypothe s is * « • « * • » . * « * . « • 131 V II. S U M M A R Y -................................. 135 LIST OF REFERENCES............................. 139 APPENDIX A ................................................ 346 APPENDIX B ...................................................................... I 69 lv Table 1* LIST OF TABLES Page E xperim ental C onditions «••••• •» •• *»1 v LIST OF FIGURES Figure page I* Schem atic diagram o f th e re p ro d u c tiv e cycle* ••••• ............ ••••• 10 2* H ypothesized i n t e r - r e l a t i o n s o f d riv e c e n te rs i n a s t a t i c s t a t e ; i* e* w ith o u t d i f f e r e n t i a l e x te r n a l stim u la tio n * • • • 29 3* Wooden model o f a m ale s tic k le b a c k and th e wooden model b la n k used fo r p r e - a d a p ta tio n • • • * • • • • • • • • « • • lj.8 l{ .* Mean nb ite a M p e r m inute f o r groups R M and M M over te n days** «• * • • • « • • • • 60 5* Mean no r le n ta tlo n s n p e r m inute f o r groups R M and M M over te n days* * * « * * o » * 6$ 6 * Median * z ig -z a g sn p e r m inute d ire c te d a t th e r e a l male over te n d ay s. • • • • * • 69 7* Change i n th e number of "z ig -z a g s* d ire c te d a t th e fem ale stim u lu s d u rin g sex t e s t s f o r a l l th re e groups over t e n days * * • 7l|- 8 * G raphic d a ta f o r f i s h number 8ID and number 13D-2, group RM • • • « « * • • « 80 9* G raphic d a ta f o r f i s h number 9BP-5 and number 12DP-5* group RM. • • • • • • • • 8lj. 10* G raphic d a ta f o r f i s h number IflD and number 7U>» group R M * * * * . . . * . * 88 11* G raphic d a ta f o r f i s h number *>DP-5 and number TDf group RM. * • • • • • • • • * 91 12* G raphic d a ta f o r f i s h number 10D and number 3®* group M M . • • • • • • • • • • 95 vi F ig u re Page 13. Graphio d a ta f o r f i s h number $d and number 11D-2, group M U .................................... 98 lit* Graphic d a ta f o r f i s h number 1 3 L D and number 63D ® group M M . . . . . . * * * . 101 15. G raphic d a ta f o r f i s h number 9D and number l*D, group M M • • » » • • • * « • 101}. 16 • Graphio d a ta f o r f i s h number 3D and number 2D-2, group 108 1 7 . G raphic d a ta f o r f is h number 2D and number 1DF-5* group C* • . ® ....................... 111 18 ♦ Graphic d a ta f o r f i s h number lljD-2 and number 13DP-5* group C .................................... 113 19. Graphic d a ta f a r f i s h number ID and number 12D, group G « « •• « • • • • * . 116 vii CHAPTER I INTRODUCTION R ecent re s e a rc h w ith b ird s (Hlnde# 19^+a, b ; I 9 6I 1 Melzack# 196l ) h a s c le a r ly dem onstrated waning o f a r e p e a te d ly e l lo l te d fix e d a c tio n p a tt e r n (FAP)** These PAPa were e l i c i t e d by re p e a te d p re s e n ta tio n s o f a stim u lu s th a t norm ally e l i c i t s t h a t PAP® Many in s ta n c e s have been re p o rte d of s tim u lu s -s p e c ific w aning o f sim pler# r e f le x iv e resp o n ses* Such phenomena are commonly c a lle d h a b itu a tio n when th e y meet c e r t a i n c r i t e r i a w hich d i f f e r e n t i a t e b e tw een h a b itu a tio n # fa tig u e # and sen so ry ad a p ta tio n # The e x te n s io n o f th e fin d in g s c ite d above to o th e r s itu a tio n s In w hich r e p e t i t i o n o f a s p e c if ic stim u lu s r e s u lts I n the w aning of a PAP re q u ir e s a s i t u a t i o n and an organism th a t i s amenable to a r e l a t i v e l y com plete a n a ly s is of I t s b e h a v io r I n th e e x p e rim e n ta l s i t u a t i o n and i s e a s ily m ain tain ed i n th e la b o ra to ry * Such a sp e c ie s l a the th re e -s p ln e d s tic k le b a c k (P a s te r os te u s ^"The fix e d a c tio n p a tte r n I s d efin ed as a se quence o f co o rd in ated m otor a c tio n s th a t ap p ears w ith o u t th e anim al h av in g to le a r n i t by th e u su a l le a rn in g p ro cess* The anim al can perform i t w ith o u t p re v io u s e x e rc is e and w ith o u t h av in g seen an o th er s p e c ie s member do i t * " (H ess, 19 6 2, p* 173*) aculoatua L*) about which a great deal ia known regarding both stim u li and FAPs as w ell as other behaviors m anifest during the reproductive cyole* I t i s the purpose of t h is research to in v e stig a te the long-term response changes brought about by prolonged p resen tation o f a stim ulus th at normally i s s u ffic ie n t to e l i c i t a FAP* The FAPs to be e lic it e d repeated ly w ill be components o f aggressive behavior* Several FAPs s p e c ific to aggression w ill be measured* On the b a sis of previous research and p ilo t work in the U n iv ersity o f Southern C aliforn ia laboratory, i t was expected th at aggressive respon siven ess would wane upon repeated e li c it a t io n of aggressive FAPs* The confirm ation of th is exp ectation was sought w ith in a m ethodological framework which allow ed in v e s t i gation o f the Independent v a ria tio n of a number o f be havior p attern s that normally occur in response to stim u li other than th ose that were used to e l i c i t aggres sion* Seme o f these behaviors ere termed "displacement a c tiv itie s " (l* e * , behaviors occurring "out of co n tex t" )* R ecently a theory of d rive in te r a c tio n has been presented th at appears to exp lain the occurrence o f d is placement a c t iv itie s * This th eory, c a lle d the " d isin - h ib ltlo n h yp oth esis," when combined with the in h ib itio n th at should r e s u lt as a concomitant of h a b itu a tio n , leads to s e v e ra l s p e c if ic p re d ic te d changes i n b eh av io r th a t can be ev a lu a te d w ith in th e framework of t h i s research * The s e le c tio n o f the th re e -s p in e d s tic k le b a c k was d ic ta te d by th e w ealth o f d e s c rip tiv e ln fo n a a tlo n re g a rd in g I t s b eh av io r p a tte r n s I n th e v a rio u s s ta g e s o f re p ro duction* A summary d e s c rip tio n o f th e re p ro d u c tiv e be h a v io r i s a n ecessary p re r e q u is ite to th e development of t h i s study* CHAPTER XX B A C K G R O U N D The Reproductive Behavior o f the Three-Opined stick leb a ck The re p ro d u c tiv e c y c le of th e th re o -sp in e d s t i c k l e back may be d iv id ed in to th re e p h a se s; ( 1) n e s t-c o n s tru c - tlo n * (2) s e x u a l, and (3) p a re n ta l* These phases and t h e i r accompanying b eh av io r p a tte r n s are d eso rlb ed i n a number o f lengthy* v ery com plete w orks. The m ost a v a il a b le of th e se works are th o se o f van I e r s e l ( 1953) and S e v e n ste r (1961)* The H e st-C o n stru c tlo n Phase I n n a tu r a l su rroundings a male w ill* a f t e r m ig ra tio n in to r e l a t i v e l y shallow f r e s h w ater* s e le c t an are a t h a t he w i l l defend and I n rthlch he w i ll b u ild h is n e s t. T his a re a i s h is " t e r r i t o r y " and i t i s u s u a lly dem arcated by b o u n d aries of p la n ts and o th e r n a tu r a l re fe re n c e p o in ts . D uring t h i s p e rio d th e m ale b eg in s to tu r n re d - orange u nder h is c h in and h is eye tu rn s a b r i l l i a n t b lu e . T his c o lo r change i s the b eg in n in g o f " n u p tia l c o lo ra t io n ." T e r r i t o r i a l i t y h as been stu d ie d i n some d e t a i l in th e la b o ra to ry by van I e r s e l (1958)* l a a. te n g a llo n k tank a male stick leb a ck w i l l normally adopt th e e n tir e tank aa h is te rrito ry * Nest b u ild in g s ta r t s w ith the male stic k le b a c k d iggin g a hole In the sandy substrate* A com plete re sponse o f the behavior la b eled "sand-digging”* ’ i s composed of the f is h p ick in g up a mouthful o f sand, swimming a short d istan ce away w ith back arched, and s p it t in g i t out* Sand d iggin g w i l l continue u n t il a shallow p it i s pro duced* Next, th e fis h w i l l p ick up sm all b it s o f algae or oth er plant m a te ria l* carry them In h is mouth and d ep o sit them In the n est p it* flftis a c t iv it y la c a lle d " collectin g*" A th ird a c tiv ity * *g lu in g ,” occurs along with c o lle c tin g * b lu in g * c o n sists o f the f is h p ressin g down h is ren al opening a g a in st the now-forming n e st and se c r e tin g a s tic k y m ateria l from h is kidneys* The shape of th e n est i s brought about by a s e r ie s of movements* the most conspicuous o f which I s a p a r tia l boring in to an entrance o f the nest* This Is o a lle d "boring*" A lso prominent during th is sta g e i s "pushing*" “Pushing” I s perpendicular or nearly perpendicular movement o f the body o f the f i s h w ith the snout pushing again st the top or s id e s of the nest* This behavior i s an In te g r a l part ^The FAPs mentioned in th is and subsequent chapters and s e t o ff by quotation marks are l i s t e d w ith th eir ob servation al id e n tific a tio n c r it e r ia in Appendix B* of th e n e s t shaping and form ing p ro c e s s . The end r e s u l t of th e above d e sc rib e d b eh av io rs i s th e fo rm a tio n of a mound of m a te ria ls p laced I n a p i t w hich has an en tran ce* The f i n a l b eh av io r p a tte r n o f th e n e s t- c o n s tr u c tlo n phase occurs when th e f i s h s t a r t s a b o rin g ty p e of movement a t the e n tra n c e o f th e n est* b u t co n tin u es squirm ing a l l th e way th ro u g h th e n e st* form ing a tu n n el* T his b eh av io r i s c a lle d "creep in g -th ro u g h * and th e f i r s t In sta n c e o f t h i s "c reep in g -th ro u g h " m arks th e t r a n s i t i o n from th e n e s t- c o n s tru e tio n phase to th e se x u a l phase* The S exual Phase The m ost prom inent c h a r a c te r is tic o f th e sex u a l phase i s th e w illin g n e s s o f th e male fis h * now w ith a f u n c tio n a l n e st* to c o u rt r i p e fem ales* D uring th e t r a n s i t i o n from n e s t- c o n s tr u c tlo n to th e se x u a l phase* th e n u p tia l c o lo rs become g r e a tly enhanced* alth o u g h th e im p ressio n of a b so lu te t r a n s i t i o n betw een phases i s some what d ecep tiv e* I n th e se x u a l phase th e m ale s tic k le b a c k s t i l l engages in a c e r ta in amount of n e s t b u ild in g a c t i v i t y of th e ty p es b r ie f ly d e sc rib e d above* and d u rin g th e n e s t-c o n s tr u o tio n phase th e re i s some evidence of c o u rtin g b e h a v io rs (van I e r s e l* 1953)* I t i s the marked q u a n tita tiv e change I n th e se b e h a v io rs th a t should be s tre s s e d * W ithin th e se x u a l phase* th e re a re two sub phases* D uring th e se x u a l phase th e m ale w il l c o u rt any fem ale th at happens In to or near h is te r r ito r y and he w i l l attempt to induce her to spawn* This s itu a tio n c o n s titu te s the sub-phase c a lle d "courtship*" The most n o ticea b le behavior# and whose frequency of occurrence i s used to in dex the strength o f sexu al responsiveness# i s a "zig-zag" movement toward the female* I f th e female i s recep tiv e# she w i l l assume a head up# t a l l down p o stu re and o rien t toward th e male# who# in turn# w ill sto p h is " zig-zags" and "lead" the fem ale toward the n e st and "show" her th e n est entrance* This "showing" by th e male c o n s is ts o f the male p o in tin g h is snout in to the n est-op en in g and turning over to allow the fem ale to e n te r the n e st* "Leading" and "showing" become more fr e quent# are recogn ized e a s ily when the cou rtin g tendency i s stron g and the fem ale i s re c e p tiv e and makes th e cor r e c t respon ses to th e m ale's "zig-zag" dance* The rea d in ess to court th e female remains a t a high but stea d y le v e l u n t il th e male a c tu a lly i s su c c e ssfu l in e n tic in g the female in to the n e st th ere she spawns* S exu al resp on siven ess w ill th en disappear fo r many m inutes (up t o one hour) and then retu rn s again* This p ro cess can be repeated for up to perhaps seven su cce ss f u l f e r t il iz a t io n s ; each tim e sexual resp o n siv en ess wanes i t recovers more slow ly* The presence o f eggs i n the n e st i s resp o n sib le f o r the tr a n s itio n from the sexu al to th e p a r e n ta l phase (S e v e n ste r, 1961)* The p re se n ce o f a n o th e r male o r n o n -g rav ld fem ale i n th e r e s id e n t m a le 's t e r r i t o r y d u rin g th e se x u a l phase fre q u e n tly r e s u l t s i n In te n s e a g g re ssiv e b e h a v io r, which i s th e second sub-phase d u rin g th e se x u a l phase • Aggres siv e b eh a v io r i s c e n tr a l to th e p re s e n t stu d y and i s d e sc rib e d i n a s e p a ra te s e c tio n b eg in n in g on page 9 * The P a re n ta l Phase T his re s e a rc h w il l d e a l w ith e v e n ts o c c u rrin g d u rin g th e se x u a l p h ase, hence th e f i s h w i ll never be allow ed to e n te r th e p a r e n ta l phase d u rin g th e e x p e ri m ent. However, a d e s c r ip tio n o f s e v e ra l b e h a v io rs th a t occur d u rin g th e p a r e n ta l phase i s in d is p e n s ib le f o r u n d e rsta n d in g th e developm ent o f th e e th o lo g lc a l th e o rie s d isc u sse d i n t h i s p a p e r. These th e o rie s were developed to e x p la in th e occu rren ce o f 1 1 o u t o f c o n te x t1 1 b e h a v io rs such as p a r e n ta l resp o n se p a tte r n s d u rin g th e sex u al s ta g e . A fte r one o r more c lu tc h e s o f f e r t i l i z e d eggs are in th e n e s t a change o ccu rs i n th e male s tic k le b a c k 's b e h a v io r. H is n u p tia l c o lo rs fa d e . He spends a g r e a te r and g r e a te r amount o f tim e "fa n n in g 1 1 and a f t e r th re e o r fo u r days " fa m in g " has become h is dominant a c t i v i t y . At the same tim e he grad u ally becomes se x u a lly unrespon siv e to fem ale stim u li and a fte r th ree or four days w i ll n o t c o u rt a resp o n siv e fem ale* "Panning” I s th e p re dominant p a tte r n I n th is phase and I t la c h a ra c te riz e d by a p o stu re o f p o in tin g downward a t an angle o f approxi m ately 30° from h o riz o n ta l w ith th e snout aimed a t th e n e s t e n tra n c e . The p e c to ra l fin s move as i n th e manner o f swimming backwards* w h ile the t a i l moves as i f p ro p e llin g th e f i s h forw ard. The n e t r e s u l t i s t h a t th e f i s h rem ains sta tio n a ry * b u t produces a stream o f w ater which flow s over th e n e s t. A fan n in g bout may l a s t from a second to as much as two m inutes* b u t i s u s u a lly i n th e fiv e to f i f t e e n second range (S evenster* 1961* and p e rso n a l o b serv atio n )* T h is behavior has th e e f f e c t o f c re a tin g c u rre n ts th a t remove w ater th a t has become s a tu ra te d w ith carbon dioxide through m etab o lic p ro c esses w ith in th e eg g s. T his fan n in g beh av io r i s m ost ch arac t e r i s t i c o f th e p a re n ta l phase* b u t i t can a ls o ooour as a displacem ent a c tiv ity (S e v e n ste r, I 96I)* A ggressive Behavior During the Reproductive Cycle F ig u re 1 i s a diagram of th e f lu c tu a tio n s of sex u al b eh av io r (Indexed by n z ig -z a g sn in sex te s ts * d escrib ed In C hapter IV)* ag g ressiv e behavior (a s Indexed by " b ite s " and "ch arg es” d u rin g a g g ressio n te s ts * des crib e d in C hapter IV)* b u ild in g a c t i v i t i e s and "fan n in g ” d u rin g th e normal b reed in g cy cle (from Sevenster* 1961* xo F ig* 1 .—Schem atic diagram of th e re p ro d u c tiv e cycle* Long term f lu c tu a tio n s i n n e s t c o n s tru c tio n ( • • • • • • ) » I n ag g ressiv e a c ti v it y ( i n sex u al a c ti v it y ( — ---- - ) , and i n p a re n ta l b eh av io r (— — )• CT * creep in g through* F * f e r t i l i z a t i o n * H * h a tc h in g of c lu tc h of eggs (from S evenster* 1961)* LEVEL O F P C 'X N V X 'f CT TIME IN DAYS 12 p* 19)* From th is diagram i t la c le a r t h a t d u rin g the se x u a l p h a se , e*g*, from th e f i r s t creep in g -th ro u g h to f e r t i l i z a t i o n , th a t both sex and a g g ressio n m a in ta in them selves a t a h ig h and s ta b le lev el* The norm al method u se d by th e European la b o ra to rie s to in s u re the m ain te nance of t h i s le v e l i s to s tim u la te th e male f i s h w ith a b r i e f exposure to a gravid fem ale h eld i n a c le a r g la ss 2 tu b e every day or two* I t should be p o in ted o u t th a t m ale s tic k le b a c k s t h a t are l e f t alone do, on o ccasio n , spontaneously r e v e r t t o th e n e s t-c o n s tr u c tio n phase and re b u ild t h e i r n e s t o r move i t to another sp o t and s t a r t an o th er cycle* How e v e r , tr a n s i ti o n from th e n e s t-c o n s tru c tio n phase to the s e x u a l phase c h a r a c te r is tic a lly in h ib its n e s t b u ild in g * I f the n e s t i s destro y ed a f t e r being p a r t i a l l y b u i l t d u rin g th e n e s t-c o n s tru c tio n phase, th e f i s h w ill com mence b u ild in g ag ain w ith l i t t l e h e s ita tio n * However, i f the f i s h suoceeds i n creep in g -th ro u g h and th u s e n te rs th e sexual phase, rem oval o r d e s tru c tio n of th e n e s t r e s u l t s i n a c e s s a tio n of a c tiv it y w ith no n e s t r e b u ild in g fo r s e v e ra l h ours or more (G u ito n , I960)* o I t h a s been th e experien ce in th e U n iv e rsity of S o u th ern C a lifo rn ia la b o ra to ry th a t a h ig h le v e l of a g g re ssiv e n e ss (and s e x u a lity ) can be m aintained a f te r n e s t com pletion (a rougher e stim a te of th e end of th e n e s t-o o n s tru c tio n phase) f o r n in e or te n days w ith o u t any s tim u la tio n from o th e r fis h * 13 A ggressive re s p o n s iv ity i s s ta b le d u rin g th e sexual sta g e o f th e re p ro d u c tiv e c y c le ; t h a t is * agg res siv e re s p o n s iv lty f lu c tu a te s very l i t t l e a f te r th e n e s t- c o n s tru c tio n phase I s com plete and sex u al re s p o n s iv lty I s n ear asym ptote* I t w ill rem ain s ta b le u n t i l th e f i s h re tu r n s to th e n e s t-e o n s tru o tio n phase and re b u ild s or u n t i l he has v ia b le eggs i n h is n e s t and e n te rs th e p a r e n ta l phase* A ggressive behaviors demonstrated by t e r r it o r ia l male stiok leb ack s are o fte n Intense but r a r e ly f a t a l to the recip ien t* The attack s u su a lly c o n s is t of a "charge* at the intruder freq u en tly follow ed by a "bite* or follow ed by a s e r ie s o f " b ite s." This a c tio n u su a lly r e s u lts in the Intruder tak ing the e a r lie s t p o ssib le opportunity to leave the area. Sometimes the b a ttle takes p lace between r e sid e n ts o f adjoining te r r ito r ie s * These skirm ishes may be longer and q u ite in ten se w ith the two p rotagon ists going around in c ir c le s try in g to b ite a t each o th er 's t a i l and/or chases and r e tr e a ts back and fo r th across the boundary between the t e r r i to ries* Such attacks are freq u en tly d irected at fem ales as w ell* i f e ith er the female i s ready to court but the male i s not* or i f the fem ale happens in to the te r r ito r y in non-gravid condition* In any case* the two most conspicuous patterns in aggression are the "charge* and th e ttbite*** W ith t h i s background in fo rm a tio n re g a rd in g th e b eh av io r of th e s p e c ie s to be used i n t h i s re se a rc h * th e co n c ep tu al and m eth o d o lo g ical c o n s id e ra tio n s o f th e means whereby resp o n se decrem ent ( h a b itu a tio n o f a g g re ssiv e re sp o n se s) may be bro u g h t about i s co n sid ered i n the nex t ch a p te r* C H A P T E R I I I P R O B L E M H abituation, Fatigue and Sensory Adaptation A stim ulus that r e lia b ly e l i c i t s a response In an organism w ill sometimes, upon frequent su ccessiv e r e p e ti t io n s , r e s u lt in a decrement In frequency and/or magni tude of th at response* The reason fo r th is decrement may be a recep tor process change, sensory adaptation, fa tig u e or h a b itu a tio n . I t has been customary to c a l l the p rocess resp on sib le fo r the response decrement habitua tio n only in cases where there i s some evidence by which both sensory adaptation and fa tig u e may be ru led o u t. In th e process of elim in ation of the two la t t e r p o ssi b i l i t i e s i t becomes p o ssib le to ascrib e p o s itiv e des c r ip tiv e p rop erties to the phenomenon c a lle d h a b itu a tio n . Fatigue i s u su a lly considered an effector-m otor p ro cess. Hence, i f fa tig u e were resp on sib le fo r the decrement in respon siven ess observed in an experim ent, no stim ulus would be able to e l i c i t the resp on se. The decrement would not be "stim ulus s p e c ific ," but rather "response s p e c ific ," I f su ccessiv e r e p e titio n of the stim ulus r e s u lts in receptor change, such as a dim inution 16 in s e n s it i v it y , t h is dim inution would a ffe c t a l l stim u li impinging upon th a t recep tor organ, not ju st th e s p e c ific stim u li (and those very sim ila r to them) th at brought about the adaptation* The decrement would be "m odality sp e c ific " but not "stim ulus s p e c if ic •" Thus stim ulus s p e c if ic it y may be used as a c r ite r io n of h ab itu a tio n as opposed to sensory adaptation and fa tig u e* The e f fe o ts o f both sensory adaptation and fa tig u e are u su a lly considered tr a n sie n t, I*e *, the decrem ental e f f e c t s o f th ese p rocesses should d issip a te r a p id ly , whereas h a b itu a tio n , lik e many forms o f lea rn in g , per s is t s * Thus temporal p ersisten ce emerges as a major c r ite r io n o f habituation* S tu d ies o f h abitu atory re sponse decrement have only r a r e ly adhered to procedures th at would c le a r ly allow u n q u alified In terp re ta tio n as h a b itu a tio n d iffe r e n tia te d from fa tig u e and adaptation* A s e le c t iv e review o f the lite r a tu r e on habitu atory response decrement organized in p h y le tic progres sio n appears as Appendix A* In th a t review no attempt was made to organize along lin e s other than phylogeny except fo r m ethodological adequacy and an emphasis on stu d ie s th at allow d iffe r e n tia tio n between h a b itu a tio n , fa tig u e and sensory adaptation* No d iffe r e n tia tio n in s e le c tio n o f papers reviewed was made in terms o f the com plexity of the response studied* 17 H a b itu a tio n ; M ethodological and Conceptual Considerations In p re v io u s s tu d ie s of h a b ltu a to ry resp o n se d ecre ment (see Appendix A) s e v e ra l im p o rtan t m eth o d o lo g ical c o n sid e ra tio n s p re se n t them selves b o th i n term s of n eces sa ry c r i t e r i a t o c le a r ly d if f e r e n tia te h a b itu a tio n from fa tig u e and sen so ry a d a p ta tio n and i n term s of m ajor param eters known to In flu e n c e ( a t l e a s t i n some c o n d itio n s and i n some s p e c ie s ) th e r a te o r n a tu re of th e h a b itu a to ry change in re s p o n s iv ity to a stim u lu s* S tim ulus I n te n s ity end H a b itu a tio n R ate A number of s tu d ie s a t d if f e r e n t p h y le tlc le v e ls have dem onstrated th a t s tim u li a t i n t e n s i t i e s n e a re r th re sh o ld are more q u ic k ly h a b itu a te d th a n a re more i n ten se stim u li (e * g », D anish, 1921; H Inde, I95^b; O ld fie ld , 1937)* I f we assume t h a t h a b itu a tio n i s due to th e development o f n eu ral in h ib ito r y p ro c esses and th u s occurs as a p o s itiv e fu n c tio n o f th e number of t r i a l s o r le n g th of time o f stim u lu s p re s e n ta tio n n ecessary to reao h a h a b itu a tio n c r i t e r i o n , we can th en vary i n h i b i tio n by v ary in g the s tre n g th o f th e s tim u li to be h a b itu ated* 18 Long Term Temporal P ersisten ce Temporal p ersisten ce has been demonstrated in a number of ways In previous stu d ies o f h a b itu a tio n . The Importance o f such demonstrations i s to show th a t the e f f e c t s o f experience w ith the stim ulus in the production o f response decrement are not simply a m atter of fa tig u e or sensory adaptation, both of whose e f f e c t s should be tr a n sito r y as pointed out above. Previous stu d ie s o f h ab itu ation concerned w ith th is problem ty p ic a lly in te r p o la te a r e s t period e ith e r in the form o f between- stim ulus-blooks r e s t in te r v a ls or between-days recovery p erio d s. N on -fatigu e, non-sensory adaptation e f fe c t s are then demonstrated in one of two vrayat e ith e r there i s le s s than f u l l recovery o f response magnitude (or response p rob a b ility ) from day to day, or w hile i n i t i a l l y recovery may retu rn the response to the previous le v e l, subsequent h ab itu a tion i s more rapid during each su cces siv e stim ulus p resen tation period a fte r the recovery perlodo Components of a Behavior C lass In a number of stu d ie s o f h ab itu ation o f complex behaviors Indexing a c la s s of b eh aviors, i t was found th at components o f the behaviors in question did not 19 h a b itu a te a t th e sane r a t e ; th a t is * seme components d isap p eared b e fo re o th e rs (H lnde, 1961? M elzack, I 96I 3 B re a s, 1952; P re c h t, 1958; s z le p , 19&|.)* T h e re fo re , i n th e p re s e n t stu d y th e m ajor re sp o n se s th a t have been re p o rte d as b ein g components o f a g g re ssio n w ere reco rd ed i n o rd e r to t e s t th e g e n e ra lity o f t h i s fin d in g * The c e n tr a l theme of th is re s e a rc h h a s to do w ith in v e s tig a tin g th e co u rse of resp o n se decrem ent o f ag g res siv e b eh a v io rs i n th e th re e -s p ln e d s tic k le b a c k as a r e s u l t of re p e a te d ex posures to s tim u li th a t norm ally e l i c i t aggression* Beyond t h i s , how ever, th e q u e stio n o f That o th e r changes i n th e o v e r a ll b e h a v io r o f th e s tic k le b a c k m ight come about as a r e s u l t o f h a b itu a tio n of one re sp o n se p a tte r n i s an in tr ig u in g one* A r e c e n t t h e o r e ti c a l schema p u t f o r t h by s e v e ra l European e th o lo g i s t s (van X ersel and B ol, 1958; S e v e n s te r, I 96I ) , p rim a rily as an a ttem p t to e x p la in disp lacem en t a c t i v i t i e s , may be combined w ith a co n cep tio n o f an in h ib ito r y s t a t e b rought about in th e co u rse o f h a b itu a tio n to p r e d ic t some s p e c if ic a lte r a tio n s i n b eh av io r p a tte r n s o th e r th a n aggression* A d e s c rip tio n of disp lacem en t a c t i v i t i e s i s a n e c e ssa ry p r e r e q u is ite to th e developm ent o f th e t h e o r e tic a l soheme in tro d u ced above* 20 Displacement A c tiv itie s D isplacem ent has been d efin ed b o th th e o r e tic a lly and e m p iric a lly by s e v e ra l authors* The most s tr a i g h t forw ard d e f in itio n i s Tinbergen* s (19f>2)s nA d is p la c e ment a c ti v it y i s an a c tiv ity * belonging to th e ex ecu tiv e m otor p a tte r n o f an i n s t i n c t o th e r than th e I n s tln o t( s ) a c tiv a te d * ” I n s tin c t* here* r e f e r s to aggression* sex* n e s t-b u ild in g , and p a re n ta l* r a th e r than th e b roader ca te g o ry of "re p ro d u c tiv e behavior" o r "re p ro d u c tiv e i n s t i n c t . " " I n s tin c t" i s e q u iv a le n t to "d riv e " or " c e n te r" on th e th e o r e tic a l le v e l used by S ev en ster (1 9 6 1 ). Tinbergen and van I e r s e l (19^7) published an a r t ic le s p e c if ic a lly on the displacem ent rea ctio n s of the th ree-sp lned stick leb ack * Tinbergen and van Ie r se l d escrib e sev era l displacem ent a c t iv it ie s th at occur a fte r the male stick leb a ck has entered the sexual phase* Two a c t iv it ie s normally associated w ith the n e st- con stru ction phase* "pushing" and "gluing*" are c la s s if ie d freq uently as displacem ent a c t iv it ie s * A third displacem ent a c t iv it y i s "fanning*" which i s normally observed in* and appropriate to* the parental stage* (A d e ta ile d a n a ly s is of "fanning" as a displacem ent a c ti v i t y has been perform ed by S evenster* I 9 6I*) 21 These displacem ent a c t i v i t i e s c h a r a c t e r is t i c a ll y occur in a sequence s ta r tin g w ith "p u sh in g ," follo w ed hy a few seconds of "fa n n in g ," and culm inating w ith " g lu in g ." F u rth e r, i t h a s fre q u e n tly been observed t h a t th e m ale s tic k le b a c k may "bore" i n th e n e st opening d u rin g th e I n i t i a t i o n of th e displacem ent sequence and in s te a d o f " g lu in g ," the f i s h may "creep -th ro u g h # " Most o fte n , however, "pushing" and "fanning" only w i l l be se e n , w ith "fanning" fre q u e n tly o c c u rrin g alone* From th e f i r s t published re p o rts o f th e o b serv a tio n of displacem ent a c t i v i t i e s , some k in d of th w a rtin g or c o n f lic t between d riv e s a n d /o r resp o n se ten d en c ie s h as been invoked as an explanation# F o r exam ple, th e occurrence o f "sand digging** a t the t e r r i t o r i a l boundary by two m ales i s ex p lain ed by th e c o n f lic t (w ith in both m ales) of th e ten d en cies to f l e e (from th e opponent*s t e r r it o r y ) and a tta c k ( in defense of h is own t e r r it o r y ) # In th e se x u a l stag e th re e common displacem ent a c t i v i t i e s o cc u r: "p u sh in g ," " fa n n in g ," and "g lu in g " d esc rib ed above# The c o n f lic t (th w a rtin g ) s itu a tio n i n t h i s case i s e ith e r produced by the sim ultaneous ten d en cies to a tta c k and c o u rt th e fem ale, o r by the absence o f th e s tim u la tio n (fem ale f i s h ) n ec essary f o r th e e x p re ssio n o f th e dominant d riv e (sex)# In a d d itio n to some elem ent of c o n f lic t o r th w a rtin g th e re la an e x te r n a l stim u lu s component th a t I s of g r e a t im portance i n displacem ent a c t i v i t i e s * For a d isp lacem en t a c ti v i t y t o occur th e p resen ce o f some stim u lu s or stim u lu s complex r e le v a n t to i t seems to be n e c e ssa ry . F o r exam ple, in b o th "fan n in g 1 1 and th e "push in g 1 * and " g lu in g 1 * a c t i v i t i e s c h a r a c t e r i s t i c a l l y demons s t r a t e d as d isp lacem en t a c t i v i t i e s d u rin g th e se x u a l s ta g e are alm o st In v a ria b ly proceeded by a t r i p to the n e st* The n e s t , or a t l e a s t th e mound o f n e s t m a te r ia ls , i s a stim u lu s f o r both "p u sh in g 1 * and "gluing** d u rin g th e n e s t c o n s tru c tio n phase when th e s e b eh a v io rs a re con n e c te d w ith n e s t fo rm a tio n . "Fanning" i s c h a r a c te r is t i c a l l y e l i c i t e d by a n e s t e n tra n c e and a n e s t w ith eggs d u rin g th e p a r e n ta l p h a se . I n th e d isp lacem en t s i t u a t i o n some o f th ese same f a c t o r s seem to o p e ra te , but th e be h a v io rs are n o t r e le v a n t to th e se x u a l stag e* Changes i n th e freq u en cy o f d isp lacem en t and vacuum a c t i v i t i e s may be seen e i t h e r d u rin g th e adm inis t r a t i o n of some e x te rn a l s tim u lu s , such as a sex or ag g re s s io n t e s t , o r d u rin g p e r io d s , u s u a lly b e fo re and a f t e r some s tim u la tio n , when th e e x te r n a l s tim u li are only th o se p re s e n t when no m a n ip u la tio n o f th e en v iro n ment i s being made. I n th e p re s e n t stu d y changes i n d is placem ent b e h a v io r w ill be w atched f o r o nly d u rin g a g re s sio n p e rio d s w ith in th e sex u al p h ase ( i . e . , d u rin g th e 23 p e rio d s Than a g g ressio n e l i c i t i n g s tim u li are p re se n t)* E th o lo g ic a l Theoryt D riv e In te ra c tio n * d isp lacem en t ana tn e b is ln h lb ltlo n H ypothesis T ra d itio n a l th e o rie s of th e displacem ent phenome non have ce n te red around what has been c a lle d th e " s u r p lu s h y p o th e sis1 * (S evenster* 1961) which em anates from older* t r a d i t i o n a l e th o lo g ic a l th e o rie s t h a t concep tu a liz e d riv e o r m o tiv a tio n a l energy as b ein g b u i l t up and th en d isch arg ed th ro u g h fix e d a c tio n p a tte r n s (FAPs)# I f the d isch arg e through com pletion o f a be h a v io r sequence were blocked by th w a rtin g o r c o n flic t* th e m o tiv a tio n a l energy would co n tin u e to b u ild up (hence a " su rp lu s1 1 ) and th e n " s p i l l 1 1 o r "sp a rk over" and d is charge through th e motor system o f some o th e r d riv e* This kind of th e o riz in g was p o p u larize d by Lorenz (1935* 1950)* In L o re n z 's model each resp o n se com prising an " in s tin c t" has a s p e c if ic energy a sso c ia te d w ith i t i n th e form of " r e a c tio n s p e c if ic energy" f o r each fix e d a c tio n p a tte r n (PAP)* This concept o f m o tiv a tio n was superseded by T in b erg e n 's which s t i l l had a concept of s p e c if ic energy f o r th e h e ir a r c h ic a l ordered d riv e cen ters* b u t th i s energy flow ed from a source a t th e top of th e h e ira rc h y which may be conceived o f as a "g e n e ra l drive*"*’ S ev en ster ( I 96I) appears to accep t th e m o tiv a- tlonaX scheme o f van I e r s e l i n term s o f h is use of th e term " d r iv e ." nD rivett has been p laced i n the d e f in itio n a l p o s itio n of b ein g * • * • th e complex o f I n te r n a l and e x te rn a l f a c to r s le a d in g to g iv en behavior* (van I e r s e l and B ol, 1958)# Tbe d riv e s t a t e i s in fe r re d from be h av io r and i t s e x te rn a l source i s In fe rre d to th e ex te r n a l s tim u li p re se n t i n th e s i tu a tio n such as th e com pleted n e s t, a r ip e fem ale, e t c . I n te r n a l f a c to r s r e f e r to changes i n endocrine b alance and o th e r in te r n a l s t a t e s . The in te r n a l s t a t e s and changes are In fe rre d from changes in b eh av io r where th e re appears to be no change i n s tim u la tio n . T his d e f in itio n o f d riv e w ill serv e f o r th e d is c u ssio n below and the r e s t of t h i s d i s s e r ta t io n . Among th e newer c o n c e p tu a liz a tio n s of th e i n t e r r e la ti o n s o f th e d riv e -c e n te rs u n d erly in g response ten d en c ie s i s t h a t of van I e r s e l and Bol (1958) & B d S ev en ster ( 1961)1 i . e . , th e d ls ih h lb ltio n h y p o th esis* The d is in h ib ltio n h y p o th e sis assumes th a t th e re i s a l A thorough d is c u ssio n of th e use o f such m o tiv a tio n a l c o n s tru c ts and th e v alu e and dangers in h e re n t i n such u ses may be found i n Hinde* 1959* Also see Z e lg le r , 19& l<> f o r a d is c u s s io n of m o tiv a tio n a l theory* p a r tic u l a r ly as i t r e l a t e s to displacem ent be h av io r* 25 center** f o r eaoh group of r e la te d a c t i v i t i e s o r b eh av io r p a tte r n s , i . e . , a sex c e n te r , an ag g re ssio n c e n te r , and & p a re n ta l c e n te r ( i n the s tio k le b a o k )• When a given c e n te r i s s tro n g ly a c tiv a te d (u s u a lly by a com bination of in t e r n a l and e x te r n a l s tim u li, i t r e s u l t s i n th e e l i c i t a tio n of b eh av io rs r e le v a n t t o th o se s tim u li * When a male stio k le b a o k venders in to th e t e r r i t o r y of a re s id e n t male i n th e sex u al s ta g e , th is a c tiv a te s th e l a t t e r * s ag g res s io n c e n te r and th e re s id e n t male a tta c k s th e tr e s p a s s e r . The dominant a c tiv a te d c e n te r s , which w i l l d i f f e r during d if f e r e n t s ta g e s of th e re p ro d u c tiv e c y c le , i n h i b it o th e r c e n te rs , su p p ressin g th e e l i c i t a t i o n of t h e i r behavior p a tte r n s . I t should be made q u ite c le a r t h a t th e e x te r n a l 2 The term s "d riv e " and "center** need t o be d efin ed f o r c l a r i t y and c o m p arab ility to th e th e o r e tic a l back ground th a t t h i s re s e a rc h r e la te s t o . S ev en ster ( I 9 6I) i s c a r e f u l to d e fin e "d riv e " as a q u a n tita tiv e concept which i s th e combined In flu e n c e o f a l l of th e a c tiv a tin g and in h ib itin g f a c t o r s , b o th I n te r n a l and e x te r n a l, th a t a c t upon th e " c e n te r" and determ ine the occurrence o r non-occurrence of a b e h a v io r. The " c e n te r" i s conceived o f as a " c o o rd in a tin g mechanism." The " c o o rd in a tin g mechanism" i s d efin ed "by i t s c a p a c ity to produce th e co o rd in ated a c tio n o f c e r ta in organs which i s c h a ra c te r i s t i c f o r th e a c tiv ity " (S e v e n ste r, I 96I , p . 1 3 ). In so f a r as "d riv e " re p re s e n ts th e sum of a c tiv a tin g and i n h ib itin g f a c to r s t h a t a c t upon th e " c e n te r" and i t , i n tu r n , ap p a re n tly w ith o u t adding any m o tiv a tio n a l component of i t s own b u t r a th e r se rv in g to " tr a n s la te " m o tiv a tio n to behavior o f a c h a r a c te r is tic type ( i . e . , a g g re ssio n , sex , e t c .) subsummed by t h a t c e n te r, th e tv o term s may fre q u e n tly be used i n co n ju n ctio n w ith one an o th er i n the term " d r iv e -c e n te r ." s tim u li a c tin g as an e x te r n a l source o f d riv e serv e b oth an e n e rg is in g (re le a s in g ) and a d ir e c tiv e f u n c tio n . Dur in g the se x u a l sta g e b o th sex and a g g re ssio n c e n te rs have an in h ib ito r y In flu e n c e on th e p a re n ta l c e n te r* There i s a ls o a m u tu al i n h ib itio n betw een sex and a g g re ssio n such t h a t whenever sex i s a t I t s z e n ith * th e in h i b it io n from a g g re ssio n i s low and conversely* when ag g re ssio n i s paramount* I n h ib itio n from th e sex c e n te r i s low* These two d riv es* which a re both dom inant d u rin g th e se x u a l phase* wax and wane i n in te rd e p e n d e n t cy cles* D uring th e waxing o f one and th e waning of th e o th e r th e re w i l l be a p o in t a t *fiiich th e sum of i n h ib it io n ( t o t a l in h ib itio n ) * a c tin g on th e p a r e n ta l c e n te r from th e s e two d riv es* w i l l be le a s t* I f th e p a r e n ta l c e n te r I t s e l f i s s u f f i c i e n t l y a c tiv a te d th e b eh av io r a p p ro p ria te to th e p a r e n ta l phase w i l l appear* T his p o in t where th e t o t a l in h ib itio n I s l e a s t r e p re s e n ts " c o n flic t* " nC o n flic t" between sex and aggression* then* low ers th e t o t a l i n h i b i ti o n e x e rte d upon th e p a r e n ta l c e n te r to th e e x te n t t t h a t th e re a re e x te r n a l sources of d riv e p re s e n t to a c tiv a te th e p a r e n ta l oenter* and p a r e n ta l b eh a v io r may th e n appear* However* I f e i t h e r sex o r a g g re ssio n were to be s u b s ta n tia lly reduced i n s tre n g th th e o th e r d riv e would become dom inant and e x e rt a d e c is iv e i n h i b it io n upon a l l o th e r c e n te rs and th e only b eh av io r th a t would 21 be I n evidence would be t h a t a p p ro p ria te to th e dominant c e n te r ( S e v e n ste r, 1961)* Prom t h i s o u tlin e o f d riv e -c e n te r I n te r a c tio n accounting f o r d is ir ih ib itio n and th e emergence o f d is placem ent b ehavior s , a more g e n e ra l model of d riv e i n t e r a c tio n may be developed* To develop th is m odel the d l s - i n h ib itio n h y p o th esis i s m odified and extended by the p re se n t w rite r* An E x ten sio n of th e P is ln h ib ltlo n H ypothesis I t i s p o stu la te d t h a t when two ce n te rs a re sim ul tan eo u sly and eq u ally a c tiv a te d , t h e i r in h ib itio n of o th er c e n te rs i s dim inished* Hence w ith the a p p ro p ria te kind of e x te rn a l s tim u la tio n (e*g*» a n est) fo rm e rly i n h ib ite d c e n te rs (e*g*, th e p a re n ta l center) w i l l be a c tiv a te d dem onstrating a b ehavior p a tte rn t h a t i s a p p ro p ria te to th e d is in h ib ite d c e n te r and to th e ex te r n a l s tim u li a v a ila b le ( I n th is example "fan n in g " would be p re d ic te d to o c c u r)• To be a c c u ra te , i t I s n ecessary to p o s tu la te th a t not only i s th e p a re n ta l c e n te r d is in h ib ite d by c o n f lic ts between sex and ag g ressio n i n the stic k le b a c k * b u t so to o i s a n e s t-c o n s tru c tlo n c e n te r* Thus i t could be pre d ic te d w ith in th e same m odel th a t n o t only "fan n in g " would appear as a displacem ent a c t i v i t y , but so too 20 would "pushing* and " g lu in g ." T his e x p e c ta tio n h a s been confirm ed from o b serv a tio n s i n th e U n iv e rsity of S outhern C a lifo rn ia la b o ra to ry which shows th a t "pushing" and "gluing* very fre q u e n tly accompany "fan n in g " as a dis® placem ent a c t i v i t y . T his e x te n sio n of th e d ls ln h ib ltlo n h y p o th e sis, th e n , p o s tu la te s a c e n te r fo r n e s t-c o n s tr u c tio n , a c e n te r fo r a g g re ssio n , a c e n te r f o r sex and a c e n te r f o r p a re n ta l b e h a v io r. For each of th ese th e re are a number o f FAPs, some of which are U3ed to index th e r e l a t i v e s tre n g th s of th e se o e n te rs (d riv e s )* I f we use S e v e n ste r1s ( I 96I) symbolism w ith th e in c lu s io n o f n e s t-c o n s tr u c tio n , we can sym bolize th e dominance and su b serv ien ce of th ese fo u r c e n te rs a t any phase of th e re p ro d u c tiv e cy cle w ith c a p ita l l e t t e r s re p re s e n tin g dominance and lower case l e t t e r s re p re s e n t in g su b se rv ie n c e . These r e la tio n s h ip s betw een c e n te rs are i l l u s t r a t e d g ra p h ic a lly i n F ig u re 2 . The fo u r o e n te rs are sym bolized th u s ; n e s t-c o n s tru c tio n H o r n % ag g ressio n A or a; sex S or is; and p a r e n ta l P o r £ • Hence, th e n e s t-c o n s tru c tlo n phase would be sym bolized as Hasp, th e c a p lto l H re p re s e n tin g th e dominance of n e s t-c o n s tr u c tio n , th e low er case a , £ and £ re p re s e n tin g th e subserv ien ce of a g g re ssio n , sex and p a re n ta l be® h a v io rs . S im ila rly , th e sex u al phase would be nASp, and 29 Fig* 2*—H ypothesized i n t e r - r e l a t i o n s o f d riv e c e n te rs i n a s t a t i c s t a t e ; l* e * , w ithout d i f f e r e n t i a l e x te rn a l stim u la tio n * Tops n e s t-c o n s tr u c tio n phase* M iddles se x u a l phase* Bottoms p a re n ta l phase* N asp nasP I - 1 - * 31 th e p a r e n ta l phase would he n a sp . T his r e s e a rc h , as n o ted In C hapter U , was concerned w ith m an ip u latio n s and e v e n ts o c c u rrin g d u rin g th e sex u al stag e ( nASp) « D uring nASP b o th sex and a g g re ssio n are dom inant; a t t h i s tim e sex u al and ag g ressiv e b eh av io rs a re c le a r ly more e a s ily e l i c i t e d by a p p ro p ria te e x te rn a l s tim u li th a n are n e s t- c o n s tr u c tio n and p a r e n ta l behaviors* B ils dominance o f th e se two c e n te rs c a r r ie s w ith i t g re a t in h ib ito r y cap acity * Hence, d u rin g nABP, A i s v ery i n h ib ito r y to n and £* i n th e same m anner, S i s very In h ib ito r y to n and £ • A and S have s tro n g m utual in h ib ito r y r e la tio n s * However, w ith th e c e n te r h ig h ly e x c ite d by in te r n a l so u rc e s, th e e x te r n a l s tim u li p e c u lia r to one o r th e o th e r d riv e r a is e s i t s c e n te r to a d e a r l y dominant p o s itio n , i f only t r a n s i e n t l y , and hence only one c la s s of b e h a v io r, th a t a p p ro p ria te to e it h e r S o r A, i s e l i c i t e d i n th e norm al s itu a tio n * Thus, th e diagram of th e s it u a t io n o c c u rrin g d u rin g nASp i n F ig u re 2 a c tu a lly d e p lo ts th e S and A o e n te rs in c o n f li c t ; th a t i s , e q u a lly e x c ite d and m u tu ally in h ib itin g * As p o in ted out above, how ever, an e x te r n a l stim u lu s a p p ro p ria te to e ith e r sex o r a g g re ssio n w i l l e le v a te one c e n te r to a c le a r ly dominant p o s itio n and hence th e dom inant c e n te r w i ll i n h i b i t th e o th e r d rlv e -c e n te rs f o r th e d u ra tio n of th a t stim u lu s and f o r a tim e th e r e a f te r * The diagram s 32 I n F ig u re 2 r e f l e c t th e h y p o th esized I n t e r r e l a t io n s of th e fo u r d riv e - e e n te rs In a s t a t i c s t a t e , w ith o u t e x te r n a l s tim u la tio n from e i t h e r a male o r fem ale stim u lu s nor any o th e r e x te r n a l stim u lu s th a t m ight a c tiv a te one c e n te r o r another* W e see th u s f a r t h a t th e sum o f th e I n te r n a l and e x te r n a l so urces o f d riv e u s u a lly makes one c e n te r dom inant and le a d s , w ith th e d ir e c tiv e fu n c tio n o f th e e x te r n a l s tim u li, to a c le a r ly sex u a l or a g g re ssiv e be h a v io r p a tte r n d u rin g the nASp phase* However, th e case s t i l l rem ains w herein th e sum o f e x te r n a l and in t e r n a l so u rces o f d riv e f o r sex and a g g re ssio n are equal* In t h i s case th e two c e n te rs would d ir e c tly I n h ib it each o th e r and, In t u r n , t h e i r in h i b iti o n on o th e r c e n te rs would be removed* Thus b eh a v io rs a p p ro p ria te to n and £ would emerge prov id ed the p ro p e r e x te rn a l s tim u li were a v a ila b le in th e environm ent* From th e above i t can be seen th a t th e p re s e n t e x te n sio n o f th e d lB in h ib ltio n h y p o th e sis m a in ta in s t h a t : ( 1) d riv e c e n te rs a re a c tiv a te d by b o th in t e r n a l and e x te r n a l sources o f s tim u la tio n and t h a t th e e x te r n a l so u rce s of d riv e a ls o serve d ir e c tiv e fu n c tio n s , ( 2 ) dur in g th e th re e s ta g e s In th e re p ro d u c tiv e c y c le , d if f e r e n t o e n te rs a re dom inant and th e dom inant c e n te r(s ) which e x e r t a s tro n g in h ib itio n on th e o th e r c e n te r s , ( 3 ) i n 33 th e case of th e se x u a l phase we have two dominant c e n te rs * S and A, (ij.) i n th e case where 3 i s eq u a l to A th e in h ib iti o n o f n and o f £ by b o th A and s i s lessen ed * W e have d esc rib ed th e g e n e ra l r e la tio n s h ip s o f th e v a rio u s c e n te rs on each o th e r i n th e norm al co u rse of th e re p ro d u c tiv e cy cle w ith p a r ti c u l a r em phasis on th e se x u a l phase w ith i t s du al c e n te r dom inance. The r o le of c o n f lic t i n displacem ent h as a ls o been o u tlin e d * Large changes in th e r e l a t i v e s tre n g th s o f e ith e r A or S d u rin g th e se x u a l phase would d is r u p t th e norm al b eh av io r d u rin g t h i s phase* U sing as a guide t h i s e x te n s io n of th e d i s ln h ib i tl o n h y p o th e sis as i t has been o u tlin e d above» p r e d ic ts th e d ir e c tio n and course o f th e se d is ru p tio n s* I f we could g ra d u a lly r a i s e th e i n t e r n a l s tre n g th o f sex r e l a ti v e to a g g re ssio n by low erin g th e s tr e n g th o f a g g re ssio n i t could be p re d ic te d th a t* a t some p o in t i n th e g ra d u a l low ering o f aggression* an e x te r n a l stim u lu s w ith predom inantly m ale c h a r a c t e r is ti c s b ut w ith th e few fem ale c h a r a c t e r is t ic s in h e re n t i n th e f i s h shape* would p la c e our s u b je c t i n c o n f lic t betw een S and A* T hat is* th e sum of e x te r n a l and I n te r n a l sources o f d riv e* even w ith th e predom inantly male e x te r n a l so u rc e , o f b o th S and A would be equal* T his con f l i c t would r e s u l t i n th e d is ln h ib itlo n of th e n and £ c e n te rs and ty p ic a l displacem ent b eh av io r would r e s u lt* As t h i s p ro c ess o f in te r n a l re d u c tio n of d riv e le v e l of A p ro g re s s e s , the same s tim u li t h a t i n i t i a l l y e l i c i t e d a g g re ssio n in th e sex u al p h ase , th e n l a t e r w ith re d u c tio n of A p laced the f i s h i n c o n f li c t , would now e l i c i t p u re ly se x u a l behavior* That i s , th e sum o f the in te r n a l and e x te rn a l so urces o f d riv e ag ain w ith predom inantly male e x te rn a l so u rc e , would sum to an in e q u a lity w ith th e S c e n te r more h ig h ly a c tiv a te d th an th e A cen ter* F in a lly , we would have an ex p erim en tal s u b je c t th a t had e n te re d th e sex u al phase o f th e re p ro d u c tiv e c y c le , w ith a s in g le c e n te r dominant r a th e r th a n th e normal d u al c e n te r dominance* I f th e dominant c e n te r i s S as we have been d is c u s s in g , we would exp ect t h i s f i s h to r e a c t more se x u a lly to e ith e r a male o r a fem ale stim u lu s th a t a f i s h who possessed a more norm al d riv e in te r r e la tio n s h ip * To t e s t t h i s e x p e c ta tio n a means of re d u cin g ( in h ib itin g ) th e ag g ressiv e c e n te r by some in te r n a l source of I n h ib itio n i s re q u ire d . H a b itu a tio n as a Source of I n te r n a l I n h ib itio n H a b itu a tio n has fre q u e n tly been co n sid ered th e r e s u l t of th e developm ent of in h ib itio n to a c o n s ta n tly re p e a te d stim ulus* I n t h i s manner i t I s very s im ila r to ex p erim en tal e x tin c tio n and, in d eed , l i t t l e d is tin c tio n betw een h a b itu a tio n and e x tin c tio n i s made by some 3£ au th o rs ( e . g . , H ilg ard and M arquis, I 9I 4 .O; K onorskl, X9l}.8) * I n h ib itio n on a b e h a v io ra l le v e l may be d efin ed as n * • • any decrease i n th e s tre n g th of a resp o n se which i s occasioned by p o s itiv e s tim u la tio n , e i t h e r of th e resp o n se i t s e l f (a s i n e x tin c tio n ) o r o f some o th e r resp o n se (a s i n e x te r n a l i n h i b it i o n ) 1 * (K im ble, 1961)* At a g e n e ra l, th e o r e tic a l le v e l B ru n to n 's (1883) d e f in i tio n i s w orth n o tin g : By in h ib itio n we mean th e a r r e s t o f th e f u n c tio n o f the s tr u c tu r e o r o rg an , by th e a c tio n upon i t o f a n o th e r, w hile i t s power to ex ecu te th e s e fu n c tio n s i s s t i l l r e ta in e d , and can be m a n ife st as soon as th e r e s tr a in in g power i s removed* The im portance o f t h i s d e f in itio n f o r th e developm ent of th e p re s e n t t h e s i s i s tw o fo ld : ( 1) th e em phasis o f i n h ib iti o n o f fu n c tio n s (o rg an s) by o th e rs i s th e e s s e n tia l fo u n d a tio n of th e d is ln h ib itlo n h y p o th e s is , and ( 2) th e em phasis on a r r e s t of fu n c tio n by i n h ib itio n r e f e r s back to th e b e h a v io ra l d e f in itio n I n term s o f decrease of response by p o s itiv e s tim u la tio n le a d in g to a c e s s a tio n of th e fu n c tio n in g of th e mechanism by which a g iv en be h a v io r i s e l i c i t e d through a sta n d a rd stim u lu s* Because o f th e s im ila r ity o f h a b itu a tio n and e x tin c tio n , b o th i n te r n s o f th e o p e ra tio n s used to ob ta in i t ( r e p e t i ti o n o f a stim u lu s t h a t has been co n d i tio n e d to e l i c i t a resp o n se i n th e case o f e x tin c tio n , and th e r e p e t it io n o f a stim u lu s th a t i s in n a te ly capable o f e l i c i t i n g a resp o n se in th e case o f h a b itu a t i o n ) , and the e f f e c ts of a d d itio n a l o p e ra tio n s on th e b u ild -u p o f in h ib itio n , such as th e p re s e n ta tio n of an e x tra n e o u s, novel stim u lu s ( d is ln h ib itlo n ) w hich r e in s ta te s a t l e a s t some o f th e resp o n siv e n ess o f th e in h ib ite d re sp o n se , we may assume th a t th e in h ib ito r y pro cess involved i s a t l e a s t s im ila r In both e x tin c tio n and h a b itu a tio n and, f u r t h e r , b o th may be considered sources of I n te r n a l in h ib itio n * Stannary and P red iction s I t i s the th e s is o f t h i s re s e a rc h th a t re p eated p re s e n ta tio n s o f a stim u lu s which r e li a b l y e l i c i t s ag g re ssiv e resp o n ses in th e th re e -s p ln e d s tic k le b a c k w i l l , w ith re p e a te d d a ily ex p o su re s, become le s s and le s s r e li a b l e in e l i c i t i n g such behaviors* This i s termed h a b itu a tio n * F u rth e r, i t i s p re d ic te d th a t th e resp o n se decrem ent w ill p e r s i s t over days and be sum atlve, thus r e s u ltin g i n le s s and le s s re s p o n s iv lty as a fu n c tio n of days* The s e v e ra l responses com prising the c la s s of be h a v io rs c a lle d "a g g ressio n ” w ill n o t h a b itu a te a t the same r a t e . A ggressive b eh av io r has been d escrib ed in C hapter I I , and c o n s is ts o f "ch arg es” a t th e stim u lu s , u s u a lly follow ed by a " b ite * " The " b ite s " sometimes occur w ith o u t "charges” and appear to he of low er In t e n s it y than th e " h ite a ” th a t occur a t th e cu lm in atio n of a "c h a rg e ," Before i n i t i a t i o n of e ith e r " c h a rg in g ,” or " b i t i n g ,* o r b o th , th e f i s h " o r ie n ts " toward th e stim ulus* "O rie n ta tio n s" do n o t seem to be s p e c if ic to one c la s s of behaviors as opposed to ano th er; t h a t i s , an o r ie n ta tio n th a t precedes a sex u a l response appears to be no d if f e r e n t from th e o r ie n ta tio n th a t p reced es a "c h a rg e ," or a " b ite ," During th e course of h a b itu a tio n i t i s p re d ic te d th a t th e h ig h in t e n s ity "charges" w ill wane w hile " b ite s " p e r s i s t , and th e " b ite s " in tu r n w ill wane, The s u p p o sitio n u n d erly in g t h i s p re d ic tio n i s th a t th e more in te n se response w i l l have a h ig h e r th re s h o ld ; i « e , , i t w il l tak e more "d riv e " to e l i c i t th e response and as "d riv e " i s lowered during h a b itu a tio n the th re sh o ld fo r the more in te n s e response w i l l be reached f i r s t . In o th e r w ords, i n i t i a l l y the a g g re ssio n e l i c i t i n g stim u lu s a c tiv a te s th e c e n te r to such an e x te n t th a t th e th re sh o ld fo r b o th "ch arg es" and " b ite s " i s ex ceeded, As h a b itu a tio n proceeds and i f the in h ib itio n a c ts upon th e d r iv e - c e n te r, th e amount of a c tiv a tio n w ill lower slow ly to th e p o in t where i t i s s t i l l above th e th re sh o ld fo r " b ite s " b u t no lo n g er above th a t f o r "c h a rg e s," As th e p ro cess co n tin u es e v e n tu a lly th e a c tiv a tio n o f th e agg ressio n c e n te r should become so low 38 th a t n e ith e r th re sh o ld I s exceeded and th e r e s u l t t r i l l be b eh a v io ral h a b itu a tio n * Xt I s f u r th e r p re d ic te d t h a t th e i n te r n a l in h ib i tio n c re a te d by the re p e a te d exposures o f ag g re ssio n e l i c i t i n g stim u lu s p re s e n ta tio n s w ill a c t on the ag g res sio n " c e n te r 1 1 as i t la conceived o f w ith in th e g e n e ra l framework of th e d is ln h ib itlo n h y p o th e sis o f van I e r s e l and Bol ( 19£8 ) and S ev en ster (1961) i n such a manner th a t s tre n g th of sex r e la ti v e to a g g ressio n w ill g ra d u al ly s h i f t i n fa v o r of s e x u a lity . This w i l l be in d ic a te d i n two ways. F i r s t , i t i s p re d ic te d th a t d u rin g p re s e n ta tio n s of th e male stim ulus* th e amount of a g g re ssio n w i ll d e c lin e over days and th e amount of sex u al beh av io r d ire c te d tow ards the male stim u lu s w ill in c re a s e . Secondly* the g ra d u al ascendency o f sex as a r e s u l t of in h ib itio n of ag g ressio n w ill be shown by an in c re a se i n th e number o f sex u al resp o n ses towards a fem ale stim ulus* Furtherm ore* th e e x te n sio n of th e d is ln h ib itlo n h y p o th esis developed above p re d ic ts t h a t during a p erio d betw een th e beginning o f th e d e c lin e of ag g ressio n and b efo re the com plete ascendency o f sex* th e re w i ll be an In c re a se in displacem ent a c t i v i t i e s which w ill de c rease ag ain as sex u al behav io r becomes in c re a s in g ly dom inant. This emergence and re c e s s io n o f displacem ent a c ti v i t y would be In te r p re te d as in d ic a tiv e o f c o n f lic t betw een sex and ag g ressio n and th e concom itant d is in - h i b l tl o n of p a re n ta l and n e s t-c o n s tru c tio n ce n te rs* T his emergence and re c e s s io n o f displacem ent beh av io rs w i ll be watched fo r d u rin g th e d a lly p e rio d s when th e ag g re ssio n e l i c i t i n g s tim u li a re being p resen ted * CHAPTER IV M ETH O D The male th re e -s p ln e d s tic k le b a c k s u se d In t h i s stu d y w ere su b je c te d to e x p e rim e n ta tio n only when th e y were in th e sex u al phase o f th e re p ro d u c tiv e c y c le because i t i s d u rin g th is stag e t h a t the s t a b i l i t y o f a g g re ssiv e and se x u a l b eh av io r can be found (s e e F igure 2 , C hapter I I ) * A ll f is h u sed i n t h i s stu d y w ere s e le c te d from a colony o f s tic k le b a c k s se in e d from Film ore Creaky n e a r F ilm o re , C a lifo rn ia * Im m ediately upon a r r i v a l In the la b o ra to ry the m ales and fem ales w ere segregated* The fem ales were p laced in f i f t e e n or tw enty g a llo n a q u a ria which were l i b e r a l l y p la n te d w ith V a llls n e r ia and E lo d ea (b o th a re ro o te d )* Two o u ts id e air-pum p f i l t e r s provided f i l t r a t i o n and w a te r c ir c u la tio n f o r each fem ale colony tank* The males were p la c e d i n tw enty-tw o g a llo n a q u a ria w ith only sand co v erin g th e bottom o f th e tan k * In s ld e - th e -ta n k f i l t e r s p ro v id ed f i l t r a t i o n . No p la n ts were placed i n th e m ale colony tan k s* As m ales i n th e la rg e colony ta n k s began to take on n u p tia l c o lo r a tio n they were tr a n s f e r r e d I n groups o f th re e to fiv e t o te n g a llo n a q u a ria , w ith sand co v erin g th e bottom and a s in g le k o o u ts id e f i l t e r p ro v id in g f i l t r a t i o n and c ir c u la tio n of w ater* The m ales used In t h i s experim ent were ta k e n from th e s e te n g a llo n ta n k s when th e y dem onstrated a deepening of th e n u p tia l c o lo ra tio n and when th e y a ls o showed ag g re ssiv e behavior tow ards tank-m ates* Both th e m ales h eld i n th e colony tan k s and th o se a c tu a lly i n use as w e ll as th e fem ales m aintained f o r use i n sex t e s t s (see below) were k ep t on a 17 hours l i g h t , 7 h o u rs dark d iu rn a l cycle* A ll f i s h were fed tu b ife x worms and liv e b rin e shrimp* The m ales were fed a sm all q u a n tity o f one o r th e o th e r o f th ese foods each day* The fem ales were fed a much la rg e r q u a n tity each day, u s u a lly in th e form o f th re e o r fo u r sm all fe e d in g s each day* F requent fe ed in g s appeared to a id i n m ain tain in g a la rg e number of fem ales i n g ra v id c o n d itio n as was r e q u ire d fo r th e sex t e s t procedure* The m ales s e le c te d fo r use were removed from th e te n g a llo n a q u a ria and p laced in p a irs in an id e n tic a l s iz e d a q u a ria (51 x 27 x 31 cm) w ith approxim ately 6 cm o f sand and g ra v el co v erin g th e bottom* Hooted p la n ts of th e same v a r ie tie s used i n the fem ale colony tan k s were p la n te d along th e l e f t s id e and o n e -th ird th e d is ta n c e from the l e f t r e a r co rn er along th e back w all of th e a q u a ria and a ls o along th e f r o n t w all from th e l e f t fro n t corner* The p la n ts served two p u rp o ses: ( 1) to p ro v id e cover f o r h id in g and ( 2 ) to p ro v id e soma o f th e m a te ria l fo r n e s t co n stru c tio n * In a d d itio n a sm all q u a n tity of th re a d algae and spyrogyra was included i n eaoh ex perim ental ta n k as t h i s m a te ria l i s p a r tic u la r ly favored by stic k le b a c k s fo r use in n e s t co n stru c tio n * As soon as th e re was some evidence th a t n e s t c o n s tru c tio n had begun, th e subm issive male o f th e p a ir in tro d u ced to g e th e r was removed. The r i g h t hand area of th e bottom extending from fro n t to back and from th e r i g h t w all out 30 cm was covered w ith g la s s p la te s * This procedure r e s t r ic te d th e area in which th e s tic k le b a c k could b u ild h is n e s t by p re v e n tin g th e dig g in g o f a n e s t p i t In the a re a covered w ith g la ss* This procedure th u s allow ed f o r a c le a r a re a on th e r ig h t t h r e e - f i f t h s o f th e tank f o r p re s e n ta tio n of stim u li* I t has been shown th a t the a g g re s s iv ity of t e r r i t o r i a l male s tic k le b a c k s v a rie s In v e rse ly w ith the a g g r e s s io n - e lic itin g stim ulus* d istan ce from the n e s t (van I e r s e l , 19f> 8) ; th u s , by r e s t r i c t i n g the a re a in which th e male could b u ild h is n e s t and by leav in g a la rg e area c le a r fo r stim u lu s p re s e n ta tio n i t was pos s ib le to c o n tro l th e d ista n c e o f stim u lu s p re s e n ta tio n from the su b je c t* s n e st* A fter a f i s h had co n stru c te d a n e s t, i t was assumed th a t he had e n te re d the sex u al s ta g e o f th e r e p ro d u ctiv e cycle* The p resence o f a n e s t i s e a s ily v e r if ie d v is u a lly * Nest c o n s tru c tio n most fre q u e n tly occurs “ overnight® ; how ever, i t may ta k e lo n g e r, up to se v e ra l days* In t h i s experim ent i f th e re was no evidence o f a n e s t under c o n s tru c tio n by th e fo u rth day th e f i s h was r e je c te d , h is ta n k oleaned and a new p a ir o f males in tro d u ced * ^ On th e f i r s t day th a t a com plete n e s t was i n evidence th e p r e te s tin g began* P r e te s tin g The purpose of th e p r e te s tin g was to e v a lu a te th e le v e l and s t a b i l i t y of the sex d riv e . S t a b i l i t y o f r e sponse to a sex stim u lu s over days was used as an in d ic a tio n th a t th e f i s h had e n te re d th e se x u a l phase of th e re p ro d u c tiv e c y c le . E v a lu a tio n o f th e le v e l of th e sex d riv e was accom plished by a d m in is tra tio n o f stan d ard sex te s ts * The Sex T est A sta n d a rd sex t e s t c o n s is ts o f th e p re s e n ta tio n $ of a r ip e (g ra v id ) fem ale s tic k le b a c k in s id e a stan d ard siz e d c le a r g la s s o r p l a s t i c tu b e a t a p re sc rib e d and co n stan t d ista n c e from th e m a le 's n e s t area* In th is i I t h as been th e ex p erien ce in th e u n iv e r s ity of Southern C a lifo rn ia la b o ra to ry th a t approxim ately Q0% o f the m ale s tic k le b a c k s tr e a te d as o u tlin e d b u ild n e sts* re se a rc h a p l a s t i c tube* cm in s id e diam eter* was used* She tube extended down in to th e tan k a l l th e way to th e bottom of the aquaria* She fem ale stim u lu s f i s h had th e freedom to move up and down th e e n tir e h e ig h t of th e t u b e . ^ The tube c o n ta in in g th e fem ale s tic k le b a c k was p re se n te d in th e r i g h t r e a r c o m e r of th e e x p e ri m ental m ale1s tank* She sex t e s t la s te d f o r fiv e m inutes d a lly and was p re se n ted i n th e m orning of each day a t as n e a rly th e same tim e as p o ssib le * M easures tak en d u rin g th e sex t e s t s w ill be dlsoussed and d escrib ed below under th e heading ^Dependent V ariab les* n Sex t e s t s norm ally e l i c i t m ostly ag g ressiv e resp o n ses during th e f i r s t day or perhaps two a f te r a n e s t f i r s t ap p ears, b u t sex u al resp o n ses become dominant a f te r th a t* For a couple of days sex u al re s p o n s iv ity may in c re a se between days by a la rg e amount; however, th e number of sex u al resp o n ses w ill s ta b i liz e fo llo w in g th e ra p id in cre ase* S t a b i li t y was defined f o r th e purposes of t h i s re s e a rc h as any two days1 sex t e s t s which are w ith in 20 p e r c e n t o f each 2 I t I s common to r e s t r i c t the range of v e r ti c a l movement of th e stim u lu s fis h * However* th is same siz e d tube was used f o r p re s e n ta tio n o f a g g r e s s io n - e lic itin g s tim u li (see below) w ith th e male f i s h r e s t r i c t e d to only th e low er p a r t of th e tube* The fem ales i n t h i s sex t e s t were given more freedom to move v e r ti c a l l y to h elp overcome any stim u lu s g e n e ra liz a tio n th a t m ight have occurred betw een sex t e s t s and ag g re ssio n stim u la tio n a d m in istra tio n s because o f th e t e s t tube s im ila r ity * o th e r I n th e number of nzig -zag s* (see M Dependent V ari a b le s H below) toward th e fem ale stim ulus* This 20 p er ce n t fig u re was based on p rev io u s d a ta th a t In d ic a te d th a t once th e sex t e s t r e s u l t s are w ith in 20 p e r c e n t of each o th e r th e re i s no g re a te r v a r i a b i l i t y th e re a fte r* When the c r it e r i o n of s t a b i l i t y was reached* th e e x p e ri m ental tre a tm e n ts began* However* th e stan d ard sex t e s t was ad m in istered to each f i s h each day throughout th e d u ra tio n of th e experim ent. The sex te s t* then* provided d a ta f o r a p re -e x p e rim e n ta l c r it e r io n of response s t a b i l i t y and f o r th e e v a lu a tio n of f lu c tu a tio n s in s e x u a lity throughout th e d u ra tio n of th e experim ent* E xperim ental Treatm ents The e f f e c ts o f d a lly exposures o f norm ally aggres s i o n - e li c it in g s tim u li were ev a lu ated th ro u g i th e use of th re e ex p erim en tal groups* Two of the th re e groups r e ceived d a ily p re s e n ta tio n s o f s tim u li th a t i n i t i a l l y e l i c i t e d a g g re ssiv e responses* These two s tim u li d iffe re d in t h e i r a b i l i t y to e l i c i t ag g ressio n ; hence stim u lu s i n te n s ity d iffe re n c e s i n r a te and/or occurrence of h a b itu a tio n could be ev a lu a te d between th ese two groups* The th ir d group re c e iv e d no a g g r e s s io n - e lic itin g stim u latio n * th u s allow ing a com parison o f changes i n s e x u a lity to the sex t e s t s tim u li betw een th e two c o n d itio n s (a g g re ssio n stim u la te d g roups, 2 , and no ag g ressio n s tim u la tio n )* These com parisons were designed to t e s t th e d is in h lb l- tlo n h y p o th e sis p re d ic tio n s w ith reg ard to changes In the sex d riv e concom itant w ith changes i n a g g re ssio n brought about by th e experim ental treatm en ts* A ggression E l i c i t i n g S tim u la tio n Groups S tim ulus c h a r a c te r is tic s t h a t e l i c i t ag g re ssio n i n male t e r r i t o r i a l s tic k le b a c k s are oval shape, and e s p e c ia lly , re d c o lo r on th e a n te r io r , v e n tr a l su rfa c e (T lnbergan, 19*>1)* These s tim u li, when combined I n a stim u lu s c o n fig u ra tio n c o n ta in in g f i s h - l i k e a s p e c ts , w ill e a s ily e l i c i t aggression* These a ttr ib u te s were combined In a wooden r e p re s e n ta tio n of a male s t i c k l e back in n u p tia l c o lo ra tio n * This model was a tta c h e d to a motor w hich ro ta te d I t in a c ir c u la r p a th , 17•!? cm i n d ia m e te r, making one re v o lu tio n every s ix seconds* In th e course of p i l o t work f o r t h i s re s e a rc h i t was disco v ered th a t male s tic k le b a c k s fre q u e n tly show ’ • f r ig h t” resp o n ses to th e model as i t r o ta te s through th e w ater* This ”f r i g h t ” response tak es th e form o f the f i s h swimming r a p id ly to a p la n t p ro te c te d co rn er and rem aining m o tio n less fo r some time* O ccasio n ally th e s tic k le b a c k overcomes h is f r i g h t and w ill th e n approach and a tta c k th e model* More fre q u e n tly , however, he w ill rem ain m o tio n le ss I n h is p ro te c te d c o rn e r u n t i l th e model i s rem oved. O ther c a s u a l o b se rv a tio n s gave th e im p ressio n t h a t t h i s " f r ig h t" was n o t as pronounced when th e f i s h was p re se n te d a s ta tio n a r y m odel. H ence, a p r e - a d a p ta tio n procedure was used to a c c lim a tiz e th e male s tic k le b a c k to b o th th e p la c in g o f th e model I n th e ta n k and to i t s movement. T his was accom plished by making a wooden "b lan k model" o f a re c ta n g u la r p iece o f wood, p a in te d th e same grey c o lo r as th e background c o lo r o f th e m odel. B oth th e model and th e "b la n k model" are shown i n F ig u re 3 . The " b la n k model" was p lace d in th e s tic k le b a c k 's ta n k f o r one h o u r, b e in g removed and r e p la c e d every f i f t e e n m in u te s. This p rocedure was r e p eated fo r a second one hour p e rio d and a th ir d p e rio d o f 30 m inutes c o n s is tin g o f th e same "b lan k model" In tro d u ced and r o ta te d a t th e same speed as th e model would b e . Yfhenever p o s s ib le each o f th e s e se s s io n s was ad m in iste red on th re e su c c e ssiv e days p r io r to th e f i s h a tta in in g th e sex u al s t a b i l i t y c r i t e r i o n . However, i n s e v e ra l c a se s th e s tic k le b a c k would re a c h c r i t e r i o n e a r l i e r th a n th e ex p erim en ter had a n tic ip a te d and th e two l a s t p r e - a d a p ta tio n p e rio d s would be p re se n te d on th e same d a y . Tw enty-four h o u rs alw ays elap se d betw een th e end o f th e p re -a d a p ta tio n procedure and th e f i r s t model p r e s e n ta tio n . MODEL BLANK MODEL MALE BLUE P m i Fig* 3 .*— Wooden model o f a male s tic k le b a c k and th e wooden model blank used f o r p re -a d a p ta tio n * The second le v e l o f th e a g g r e s a lo n -e lic itin g s tim u lu s , and the one which e l i c i t e d th e g r e a te s t amount of a g g re ssio n , was a liv e m ale s tic k le b a c k , i n n u p tia l co lor* This stim u lu s was confined i n a c le a r p l a s t i c tube of th e same dim ensions as th e one used f o r th e sex te s ts * The male stim u lu s f i s h was confined to th e low er te n cm o f th e tube by a c le a r p l a s t i c p lu g which was i n s e rte d from above* The stim u lu s tube was suspended i n th e ta n k such th a t th e stim u lu s appeared a t th e same d ista n c e from th e r ig h t hand sid e o f th e a q u a ria as th e c e n te r o f th e circum ference o f th e r o ta tio n a l p a th of th e model* T his d is ta n c e was approxim ately 20 cm from th e r i g h t hand sid e o f th e ta n k , e q u id is ta n t from th e f r o n t and back w alls* Both groups, th e one re c e iv in g s tim u la tio n from the model (model male group) and th e one re c e iv in g s tim u la tio n from the r e a l male f i s h ( r e a l male group) were p re se n te d w ith th e a p p ro p ria te a g g r e s s io n - e lic itin g stim u lu s fo r 1 5? m inutes each day f o r te n co n secu tiv e days, s t a r t i n g on th e afte rn o o n of th e day th a t th e f i s h reached th e s t a b i l i t y c r i t e r i o n i n th e morning sex te s ts * The s tim u la tio n p e rio d follow ed th e d a ily sex t e s t by a t l e a s t one hour and was given a t as n e a rly th e same tim e each day as p o ssib le * The m easures ta k e n during the s tim u la tio n p e rio d s were th e same as th o se ta k e n d u rin g sex t e s t s and are d iscu ssed below under th e heading nDependent V a ria b le s •* Non-S tln n ila tlo n Group A th ir d group re c e iv e d no s tim u la tio n o th e r th an th e d a lly fiv e m inute sex t e s t each m orning. The sex t e s t s were continued fo r te n days a f te r th e f i s h had made th e s t a b i l i t y c r ite r io n * Thus* a l l groups re c e iv e d the same trea tm en t i n term s of sex t e s t s fo r th e same number of days* However* t h i s n o n -stim u la tio n c o n tro l group did n o t re c e iv e any ag g ressio n -e 11 c itin g s tim u la tio n durin g th e d u ra tio n of th e ex p erim en tal period* From t h i s group ohanges i n s e x u a lity could be In fe rre d from f i s h th a t were never p re se n te d w ith a g g r e s s io n - e lic itin g stim u li* T his group served as a b a s e lin e upon which to c o n tra s t th e two s tim u la tio n groups In term s of changes in se x u a l i t y over th e course of th e experim ent* P o st-T e sts S u b jects i n group C (n o n -stirn u lated c o n tro l group) were ad m in istered a f i f t e e n m inute ag g ressio n s tim u la tio n p erio d w ith a r e a l male in the tu b e on the a fte rn o o n of day ten* The purpose o f t h i s was to d e te c t any ohanges i n ag g ressio n th a t m ight occur o nly because o f th e te n day p erio d of the experim ent r a th e r than to the d a lly SI a d m in is tra tio n s of a g g re ssio n e l i c i t i n g s tim u li* T h is t e s t would he compared to group RM ( r e a l m ale group) on day one* S u b je c ts in group M M (model male group) were ad m in is te re d a f i f t e e n m inute a g g re s s lo n -s tim u la tio n p e rio d w ith a r e a l male in th e tube on th e a fte rn o o n of day eleven* The purpose o f t h i s t e s t was to determ ine I f th e re was any d im in u tio n In re s p o n a lv lty to a r e a l male as a r e s u l t o f th e h a b itu a tio n ex p erien ce w ith th e model* T his I s e s s e n tia lly a t e s t f o r stim u lu s s p e c i f ic i t y i n h a b itu a tio n * The a p p ro p ria te com parison i s w ith group R M on i t s f i r s t day o f r e a l male stim u la tio n * A summary of th e ex p erim en tal c o n d itio n s and th e stim u lu s a p p lic a tio n s appears i n Table 1* TABLE 1 EXPERIMENTAL CONDITIONS Group P r e te s t Experim ent P o st T est R eal Male Sex t e s t s Sex t e s t s / r e a l m ale ---- (RM) Model Male Sex t e s t s Sex te s ts /m o d e l m ale R eal male (M M ) C o n tro l Sex t e s t s Sex t e s t s / — - R eal m ale (C) Subject Assignment When a f i s h b u i l t a n e s t and h is subm issive p a rtn e r was rem oved, th e ex p erim en ter s e le c te d from a paper bag one o f th re e i d e n t i c a l t i l e s , each having a group d e s ig n a tio n in s c rib e d upon I t* The s u b je c t was assig n ed to th e group whose in s c r ip tio n was on th e t i l e s e le c te d . Toward th e end o f th e experim ent when one group was com pleted (N - 8 ) th e t i l e rem ained in th e bag but was re p la c e d i f s e le c te d and an o th er t i l e drawn. This method i s n o t s t r i c t l y random b u t ap p ears to a s s ig n s u b je c ts to groups w ith o u t b i a s .3 E ig h t f i s h com prised the sam ple fo r ea ch group* Dependent V a ria b le s C e n tra l to th e approach s e le c te d f o r t h i s re s e a rc h i s th e u se of th e w ealth of e m p iric a l d e s c rip tio n s o f re sp o n se s and b eh av io r p a tte r n s observed d u rin g th e r e p ro d u c tiv e cy cle o f th e th re e -s p in e d s tic k le b a c k . The p re s e n t re s e a rc h was designed around as r e l a t i v e l y com p le te a d e s c r ip tio n of b e h a v io r as was p o s s ib le w ith in ^Five f i s h were r e je c te d from the stu d y a f te r assignm ent to groups on th e b a s is of not resp o n d in g to the m odel even a f t e r p re -a d a p ta tio n . One f i s h managed to g e t in to h is f i l t e r o v e rn ig h t and th e r e a f te r developed a d is e a s e th a t proved f a t a l . T his su b je c t was from group RM. 53 the lim ita tio n s o f th e te c h n ic a l a b i l i t i e s of th e re c o rd in g system and th e o b se rv a tio n a l a b i l i t y of th e e x p e ri m enter. D e sc rip tio n s o f s tic k le b a c k b eh av io r a re ty p ic a l ly grouped as b eh av io rs e x p re ssin g v a rio u s d riv es* F or each d riv e th e re are se v e ra l ty p ic a l and p re v a le n t be h a v io rs . These have been term ed "d riv e in d ic a to r v a r i a b le s ." D rive In d ic a to r V a ria b le s On th e b a s is o f th e e th o lo g is t* s background of re se a rc h and o b se rv a tio n s th ey have s e le c te d one o r two v a ria b le s th a t In d ic a te and d e fin e a d r iv e . Thus "ag g re ssio n " i s d efin ed i n term s o f th e number o f "bitjes" and " c h a r g e s "Sex" i s d efin ed i n term s o f th e number of "z ig -z a g s" toward a fem ale stim u lu s i n a g la s s or p l a s t ic tu b e . "Z ig -zag s" a r e , by e th o lo g lc a l a n a ly s is , a c tu a lly a com bination of a g g ressio n and " le a d in g " (a sexual b e h a v io r), and hence n o t a p u re ly sexual b e h a v io r. However, S ev en ster ( I 9 6I ) showed th a t th e re i s a h ig h c o r r e la tio n betw een "z ig -z a g s" (which are very e a s ily recognized and counted resp o n se) and "lea d in g " (which i s le s s e a s ily re c o g n iz e d ), and th e re fo re th e " z ig -z a g " i s - See Appendix B f o r a d e s c rip tio n o f th e se in d ic a to r v a r ia b le s and t h e i r o b se rv a tio n re c o g n itio n c r i t e r i a . S t used as th e prim ary in d ic a to r o f s e x u a lity and se x u a l r e s p o n s lv lty . N e s t-c o n s tru c tlo n b e h a v io rs m ost f r e q u en tly encountered d u rin g th e sex u al phase are "p u sh in g * 1 and "g lu in g * 1 * The p a r e n ta l d riv e in d ic a to r v a r ia b le th a t i s u n iv e r s a lly agreed upon i s "fanning** F or each o f th e fo u r m ajor d riv e s (n e s t-o o n s tru o - t io n , a g g re s s io n , sex and p a re n ta l) th e re i s a l i s t o f f re q u e n tly and le s s fre q u e n tly observed b eh a v io rs w hich were reco rd ed d u rin g th e experim ent* D e s c rip tio n and o b se rv a tio n re c o g n itio n c r i t e r i a o f th e s e appear as Appendix B« X * S T est-co n stru ctio n 1* Sand p ic k in g 2 . Pushing* 3* B oring fy -* G luing* 5* M a te ria l t e s ti n g 6 . C reeping th ro u g h A ggression 1* Charges* 2 * B ite s* 3 . Sex T h re a ts 1 * Z ig -zag s* 2 * Leading* 55 IV. P a re n ta l 1 . Panning* V. Other I . Orient, a t Iona toward th e stim u lu s* The l a s t "behavior m entioned above ( o r ie n ta tio n s toward the stim u lu s) was in clu d ed f o r two re a s o n s . F i r s t , o rie n tin g may be considered an in d ic a tio n of a tte n tio n to th e stim u lu s ( v ig ila n c e ) . I t m ight be ex p ected th a t durin g th e co u rse of h a b itu a tio n th e f i s h m ight in h i b it ag g ressiv e re sp o n se s, b u t co n tin u e to o r ie n t a t l e a s t f o r a w h ile , so th a t a g g re ssio n and o r ie n ta tio n m ight show decrem ental r a te s t h a t are d i f f e r e n t. Secondly, NSlzack ( I 96I ) showed th a t w hile b ird s m ight cease nf e a r responses* to a model o f a p re d a to r, they d id n o t sto p o r ie n tin g . The o r ie n ta tio n s toward the stim u lu s m ust, by t h e i r d e f in itio n (see Appendix B) c o r r e la te w ith s e v e ra l o th e r m easures u sed , p a r tic u la r ly w ith nz ig -z a g a ,n ncharges" and " b i t e s .n R ecording and Measurement of Dependent V a ria b le s A ll th e behaviors l i s te d above (and d escrib ed i n Appendix B) were recorded d u rin g b oth th e fiv e m inute m orning sex t e s t s and d u rin g th e f i f t e e n m inute a fte rn o o n p re s e n ta tio n s of th e a g g r e s s io n - e lic itin g s tim u li. Those w ith s u p r a - s c r ip t a s te r is k s were reco rd ed by th e e x p e ri m enter d e p re ssin g a b u tto n on a sm all hand console* The b u tto n s $ upon d e p re ssio n , d e fle c te d a 1*1 /2 v o lt s ig n a l magnet w ith in k -w ri te r* R elease o f th e b u tto n re tu rn e d th e pen to th e r e s t i n g p o sitio n * The in k -w ritin g pens w rote on paper p assed beneath them a t a c o n s ta n t speed o f 11* 1 ) . cm p er m inute* A s in g le " b lip " on one of seven channels served to r e g i s t e r a l l resp o n ses ex cep t "fa n n in g * 1 f o r w hich d u ra tio n was reco rd ed by d e p re ssin g th e pen b u tto n f o r th e d u ra tio n of th e "fan n in g b o u t* ” Thus, b oth frequency and d u ra tio n were reco rd ed f o r "fanning*" O ther b eh av io rs were noted e i th e r by w ritin g d ir e c tly on the moving re c o rd paper o r by making n o te s on a sh e e t of p ap er a tta c h e d to th e hand co n so le t h a t co n tain ed th e b u tto n s* CHAPITER V RESULTS The r e s u lt s o f t h i s re s e a rc h ere p re se n te d i n two s e c tio n s . The f i r s t s e c tio n i s concerned w ith gpoup e f f e c t s , b oth w ith in - and betw een-groups* The seoond s e c tio n d e a ls w ith d a ta and c o n c lu sio n s based on i n d iv id u a l s u b je c ts and sub-groups o f s im ila r s u b je c ts w ith in th e sample* Twelve re sp o n se s were r o u tin e ly re co rd ed d u rin g th e experim ent (see C hapter IV)* However, some of th e s e re sp o n se s occurred w ith such in fre q u en cy t h a t a n a ly s is of th e freq u en cy o f t h e i r occurren ce seemed e s s e n t i a l l y m eaningless* These In fre q u e n t re sp o n se s In clu d ed " b o r - i n g ,w “c re e p in g thro ugh" and " th re a ts * " These could n o t be analyzed as group e f fe c ts * "Pushing" and "g lu in g " were fre q u e n tly found to accompany "fan n in g " a s d is p la c e ment b eh a v io r and th e r e f o r e th e most fre q u e n t d is p la c e ment a c ti v i t y was analyzed as an in d iv id u a l e f f e c t (" fa n n in g ")* "Leading" was found to be h ig h ly c o r r e la te d w ith "fan n in g " which i s u n d ersta n d ab le in s o f a r as th e re c o g n itio n c r i t e r i o n f o r t h i s re sp o n se was n o t d is tin g u is h a b le from sim p le " t r i p s to n e s t," and " t r i p s to 57 n e s t" alm ost in v a ria b ly preced ed "fa n n in g bouts*"1 Most o f th o se re sp o n se s t h a t , fo r th e re a so n s s ta te d , were n o t analy sed as group e f f e c t s were used f o r a n a ly s is o f in d iv id u a l s u b je c t d a ta and f o r sub-group groupings in th e second s e c tio n o f t h i s ch ap ter* Group E ff e c ts P r e - t e s t sex t e s t s were ad m in iste red to e v a lu a te th e le v e l and s t a b i l i t y of th e sex d riv e b e fo re th e be g in n in g of th e experim ent* The number of n z ig -z a g s 1 * made d u rin g th e se sex t e s t s by each f i s h on the two con se c u tiv e days d u rin g w hich th e d iffe re n c e betw een th e number o f " z ig -z a g s ” was w ith in 20 p e r c e n t of th e p re v io u s t e s t was combined p ro v id in g a mean number o f ” z ig zags” a t c r ite r io n * A K ru sk a l-W a llis n o n -p ar ame t r i e a n a ly s is o f v a ria n c e u sin g average ”re sp o n se s p e r m inute” f o r each f i s h y ie ld e d an in s i g n if ic a n t groups e f f e c t , H = 1*80 ( *£0 > p > *30)*^ The m edians f o r each group • i T his i s n o t to im ply t h a t " t r i p s to n e s t” and " le a d s ” are i d e n t i c a l o r v ery sim ila r* I t was n o t pos s i b l e , how ever, f o r t h i s w r ite r to d e fin e " le a d s ” i n an o b je c tiv e manner t h a t c le a r ly d i f f e r e n tia te d " le a d s ” from "n o n -le a d in g t r i p s to n e s t* ” ^K T uskal-W allis A nalyses o f V ariance were used th ro u g h o u t f o r betw een-groups o v e r a ll com parisons* S p e c ific betw een-groups com parisons were made w ith Mann- W hitney U te s ts * Both m ethods are d e sc rib e d i n d e t a i l i n S* S e i g e ^ s Non-P a rametr i c S t a t i s t i c s f o r th e Be h a v io ra l S c ie n c e s* McGraw-MjllV fiew Y ork, 1 956« 59 were* R eal Male group (RM) 2 7 .£ , Model Male group (M M ) 35*1* &nd s e x - te s ts - a lo n e - c o n tr o l (G) 30*2* From t h i s I t I s concluded th a t th e th re e groups d id n o t d i f f e r m arkedly I n t h e i r le v e l of s e x u a lity a t th e o u ts e t of th e experim ent* Two a g g re ssiv e resp o n ses* " b i t e s ’ * and " c h a rg e s ," were p re d ic te d to wane over th e course o f te n days of d a ily a g g r e s s io n - e lic itin g stim u lu s p r e s e n ta tio n . Of th e se two resp o n ses* " b ite s " was th e m ost fre q u e n tly e l i c i t e d response* F ig u re If. shows th e mean " b ite s p er m inute" f o r b o th group R M and M M over th e te n days of a g g r e s s io n - e lic itin g stim u lu s p re se n ta tio n * The graph i s p re se n te d i n term s o f f iv e g ra p h ic p o in ts fo r each day* Bach p o in t r e p re s e n ts th e mean b i t e s p e r m inute fo r th re e co n se c u tiv e m inutes* Thus, th e f i r s t p o in t f o r each day r e p re s e n ts a mean of th e f i r s t th re e m inutes* th e second p o in t* the mean o f th e second th re e m inutes* and so fo rth * A com parison of th e t o t a l number o f " b ite s " on day one betw een groups R M and M M dem o n strates a r e l i a b l e d iffe re n c e on th e i n i t i a l day p f a g g r e s s io n - e lic itin g stim u lu s p re s e n ta tio n (Mann-TShitney X T ■ 2*0* p < *001)* A p a r a l l e l a n a ly s is perform ed on th e te n th day betw een th e two groups showed t h a t th e groups were no lo n g er s i g n i f ic a n tl y d if f e r e n t alth o u g h the t e s t y ie ld e d a 60 P ig . ij.*— -Moan " b ite s ” p er m inute fo r groups R M and M M over te n days* Bach p o in t re p re s e n ts th e average of th re e su ccessiv e m inutes* 61 RM M M I l i l l 11 i t i 1 i i i i 11 i ii I i i i i l l 111 I i i i i I » i i ILi. i u i i LU. RAYS 62 T J * 17*0 (p » *065> ) w hich c o n tra in d ic a te s th e assum ption of e q u a l le v e ls of re sp o n siv e n ess between th e two groups a t th e te rm in a l p o in t of treatm en t* W ithin-groups com parisons showed th a t b o th groups decreased i n th e number of nb l t e s n over th e e x te n t o f th e experim ent* The most dram atic e f f e c t o f d a ily r e p e t i tio n s of th e a g g r e s s io n -e lic itin g stim u lu s p re s e n ta tio n s i s found comparing th e f i r s t and l a s t days of group R M * T his decrease i s s t a t i s t i c a l l y r e l i a b l e as te s te d by th e W llcoxon M atch ed -P airs, Signed Ranks t e s t , T * 0 , p « .OOl+o^ The decrease i n ag g ressio n betw een th e f i r s t and l a s t day i s l e s s dram atic fo r group M M , b u t i t i s s t i l l a r e li a b l e change as measured by the same t e s t (w ilcoxon T * 3 .0 , *025 > p > * 0 1 ). The g re a te r c a p a c ity o f the r e a l male to e l i c i t N b l t e s ,> compared to th e model in d ic a te s th a t th e model was a f a r le s s adequate stim u lu s th an was th e r e a l male* The s ig n if ic a n t d iffe re n c e s betw een the number o f • b i t e s 1 * on day one as compared to day te n f o r b o th groups shows th a t a c le a r resp o n se decrem ent e f f e c t has tak en p lace over th e te n days (F ig u re I}.)* From the f a c t th a t group M M and R M a re n o t s ig n if ic a n tly d if f e r e n t on day te n , in 3 HIMLlcoxon M atch ed -p airs, Signed-Ranks t e s t s were used throughout fo r w ith in -g ro u p s com parisons ex cep t where o therw ise noted* This method i s d escrib ed i n d e t a i l in S* S ieg el* s N on-Param etrie S t a t i s t i c s fo r the B ehavioral Sciences* New Y o rk :' McGraw-Hill, l^!>b* a d d itio n to th e g re a t d iffe re n c e between th e two groups on day one, i t can be seen t h a t th e r a te o f d ecrease and th e absolute amount o f d ecrease i n " b itin g " b eh av io r was g r e a te s t fo r group R M * I t should a lso be p o in ted out th a t comparing th e f i r s t p o in t of each day w ith th e l a s t p o in t on th e p rev io u s day shows th a t th e re i s a reco v ery o f re s p o n s iv ity to th e r e a l male over th e 23- 3/ij. hour b reak between p re se n ta tio n s* T his group e f f e c t holds f o r th e com parison betw een each p a i r o f days* Although th e number of s u b je c ts showing "ch arg es" toward the stim u lu s i n group M M i s sm all ( f iv e o f the e ig h t su b je c ts) and th e number o f "charges" evidenced by th e se fiv e f i s h i s a ls o sm all (ran g e * 2- 6, m edian * 3 ) , a n a ly s is of t h i s d a ta dem onstrates r e s u l t s c o n s is te n t w ith th e o th e r ag g ressio n m easure ( " b ite s '1 )• A between* groups a n a ly sis showed th a t group R M s u b je c ts "charged" more o fte n th a n group M M s u b je c ts (Mann-TShitney U s o*5>, p < *001) * On day one th e R M group "charged" more f r e q u e n tly th an group M M (Mann-Whitney U * 3*£» P c •0 0 1 ), b u t by day te n th e re was no d iffe re n c e nor even a tre n d toward a d iffe re n c e betw een th e groups* A w ith in -g ro u p s a n a ly s is was perform ed on b o th groups com paring th e t o t a l number o f "ch arg es" on day one and two combined w ith the to t a l number of "ch arg es" on days nine and te n combined* Group R M showed a r e lia b le d ecrease over th e course of th e experim ent (W ilcoxon T = 0 , p ■ •O O I4 .) and group M M , d e s p ite th e re d u c tio n of u sa b le s u b je c ts to 5 , a ls o shoved a r e l i a b l e d e c lin e i n th e number o f "ch arg es" (S ig n t e s t , x ■ 0 , p < *03)*^ I t i s c le a r t h a t f o r "ch arg es” as v e i l as f o r " b ite s 1 1 th e re i s a d im in u tio n i n th e number o f resp o n ses over th e e x te n t of the experim ent* I n s h o r t, th e se two ag g ressiv e resp o n ses wane w ith re p e a te d stim u lu s p re se n ta tio n s* ©1 e frequency o f " o r ie n ta tio n s " toward th e stim u lu s d u rin g th e te n days o f a g g r e s s io n - e lic itin g stim u la tio n was not expected to show any marked decrease* The r e s u l t s o b tain ed fo r t h i s measure a re p o rtra y e d g ra p h ic a lly i n F ig u re £• An a n a ly sis s im ila r to th o se used fo r th e " b ite s " and th e "charges" m easures was p e r formed on t h i s response* On day one group RM showed s ig n if ic a n tly more " o r ie n ta tio n s " toward the stim u lu s th an group M M (Mann-TiShltney X T * 5 > * 0» P * •001)* O n day te n a p a r a l l e l a n a ly s is showed th a t th e two groups d id n o t d i f f e r s ig n if ic a n tly a t th e te rm in a l p o in t of t r e a t ment ( Mann-Whitney U « 28, p » *36)* F u rth e r, a w ith in - groups a n a ly s is was perform ed f o r each group* Both ^A S ig n t e s t was used fo r t h i s t e s t because ta b le s of s ig n ific a n c e le v e ls f o r th e Wllcoxon t e s t do n o t accommodate Rs as sm all as fiv e * 65 Pig* 5* —Mean M o rie n ta tio n s " p er m inute f o r groups R M and M M over te n days* Each p o in t re p re s e n ts th e average o f th re e su cc essiv e m inutes* SAYd o / 6 9 I 9 £ It £ z / m t j t 1 1 1 11 i i r 11 i-i" i j r 1 \ r | f i 11 j- r r r i | *ii q -rt M j 11 inq 99 an a ly se s dem onstrated a s ig n if ic a n t d ecrease i n th e number o f " o r ie n ta tio n s " betw een days one and te n . Group R M showed th e g r e a te s t d ec rease as F ig u re £ > in d ic a te s (W llcoxon T = 0 , p » »00lj.). Group M M shows l e s s o f a sh a rp d e c lin e i n F ig u re *5. alth o u g h th i s to o i s a s i g n if i c a n t decrease (w ilcoxon T ® 1 , p = *035>) • The h ig h e s t t o t a l d a lly re s p o n s lv lty f o r group M M does n o t occur on day one. however# as i t does w ith group RM. W ith group M M a com parison o f day two re sp o n siv e n e ss w ith day te n y ie ld s a sm a lle r W ilcoxon T (T " 0# p a »00i{.)» I t i s c le a r from th e above an a ly se s t h a t th e two groups d i f f e r In th e number o f n o r ie n t a t Ions’* th ey make tow ard th e a g g r e s s io n - e lic itin g stim u lu s a t th e b eg in n in g of th e experim ent* Both groups show a r e l i a b l e d ecrease i n th e number o f ’ ’o r ie n ta tio n s ” th e y make d u rin g th e co u rse of th e ex p erim en t, and th e t o t a l d a ily te rm in a l le v e l of re sp o n siv e n e ss I s eq u al f o r b o th g ro u p s. The d ecrease in " o r ie n ta tio n s ” over days was n e ith e r p re d ic te d n o r e x p e c te d . I t was p re d ic te d th a ts (1) th e most ag g ressiv e re sp o n se , " c h a rg e s ." would be th e f i r s t to d e c re a se . ( 2 ) " b ite s " would be n e x t, and ( 3) " o r ie n ta tio n s " would n o t show a d e c re a s e . In th o se f i s h showing "ch arg es" i t did indeed d isa p p e a r o r re a c h a v e ry low le v e l ra p id ly * The " b i te s ." a much more fre q u e n tly e l i c i t e d b e h a v io r. 68 decreased le s s ra p id ly an d , c o n tra ry to p r e d ic tio n , " o r ie n ta tio n s ,” w hile th e y did d e c lin e over d ay s, never reach ed the low le v e l t h a t "ch arg es” and " b ite s " did* B esides " o r ie n ta tio n s ” toward th e stim u lu s and th e two indexes o f a g g re ssio n , "ch arg es” and " b i t e s , ” only one o th er recorded b eh av io r showed enough frequency a c ro ss su b je c ts to be c o n sid ered and analyzed as a group e ffe c t* In group R M a l l e ig h t f i s h "zig -zag g ed ” toward the a g g r e s s io n - e lic itin g stim u lu s a t some tim e d u rin g th e course of th e experim ent* Because th ese d a ta were so v a r ia b le , they were tr e a te d somewhat d if f e r e n tly th an the o th e r m easures re p o rte d as group e ffe c ts * The f r e quency of " z ig -z a g s” was combined i n term s of means f o r each s u b je c t f o r each b lo c k of th re e t r i a l s ju s t as th e o th e r m easures w ere, b u t th e group curves were based on the m edian of th e means r a th e r th a n on th e mean of the means* F igure 6 p re se n ts th e m edian number of " z ig -z a g s ” p er m inute* I t shows a sh arp in c re a s e between day one and day two, and then a r e tu r n w ith some d e v ia tio n s to the le v e l of th e f i r s t day* A com parison betw een day one and day two showed th a t t h i s sh arp in c re a se was s t a t i s t i c a l l y r e lia b le (W llcoxon T 8 0 , p 8 *0l6; N 8 6)* In s p e c tio n of th e d a ta re v e a le d th a t no o th e r day compared w ith day one approached sig n ific a n c e * Group M M co n tain ed 69 F ig . 6 *—Median ” z ig -z a g s” p e r m inute d ire c te d a t th e r e a l male over te n d ay s. Each p o in t re p re s e n ts th e average o f th re e su c c e ssiv e m inutes* 1/EDIAN Z IG -Z A S S /M IN . 70 6 / / 71 only one su b ject th a t showed any prominent "zig-zag" behavior, hence no group e f f e c t was present* Group RM, which receiv ed d a ily p resen tation s o f the r e a l male stim ulus fo r f i f t e e n m inutes, showed & steady d eclin e in r e sp o n siv ity to th a t stim ulus over the ten days o f the experim ental treatment* To show th a t t h is d eclin e was not sim ply due to the len gth of time th at passed between the beginning and the end of the experim ent, a f i f t e e n minute r e a l male t e s t was presented to group C (which had receiv ed only d a ily sex t e s t each morning fo r the ten days) on the afternoon o f the ten th day o f the experiment* I f time alone (te n days) were s u f f ic ie n t to account fo r the response decrease between day one and day ten fo r group R M we would expect the fifteen -m in u te-rea l-m a le t e s t on day ten fo r group C to show le s s re sp o n siv ity toward the r e a l male than the day one rea l-m a le-p resen ta tio n for group R M * A Mann- Whitney t e s t , between the day ten p o s t-te s t fo r group C and the day one real-m ale p resen ta tio n for group R M u sin g the number o f " b ite s," showed the two groups were not d iffe r e n t (U « 32, p = .3 2 )* Group M M was presented w ith a re a l male on day elev en of the experiment* The purpose of t h is p o s t - t e s t was to a sc e r ta in i f the response decrement seen with th a t group was s p e c ific to the model stim ulus or whether i t 72 was a gen eral decrement In aggression to a l l aggression - e l i c i t i n g stim u li* A Mann-Whitnay t e s t was performed comparing the number o f "bites** during th e d ay-eleven p o s t - t e s t p resen ta tio n of the r e a l male fo r group M M w ith th e number of "bites" during " H ie day-one r e a l male p resen ta tio n for group R M * No d iffer en ce was found (U » 31*!?* P > *29)* These two p o s t - t e s t comparisons show: (a) the response decrement found w ith group R M over th e te n days of the experiment was not due to the time period alone * and (b) the decrement in aggressive responses d irected to the model in group M M did not a lte r th e aggressiv e r e sp o n siv ity to the more adequate stim ulus* the r e a l male* To summarize the r e s u lts o f the d a ily f if t e e n minute a g g r e s s io n -e lic itin g stim ulus p resen tation s* i t i s clea r th a t both "b ites" and "charges" show a decrease i n frequency as a p o s itiv e fu n ctio n of th e number o f days o f stim ulus p resen ta tio n in both groups M M and R M * The frequency o f "orien tation s" toward the stim ulus shows a sim ilar* but le s s marked d eclin e over the ten days o f experim entation* The frequency of " zig-zags" toward the stim ulus by group R M increased sharply on the second day o f stim ulus p resen ta tio n and returned to a lower le v e l comparable to the count on day one* Group M M showed no "zig-zag" behavior toward the stim ulus model* The f iv e m inute sex t e s t s th a t were used to e v a lu a te th e le v e l o f s e x u a lity and th e s t a b i l i t y of th e resp o n se le v e ls of th e su b je c t were continued throughout the course o f th e experim ent* Thus, th e le v e l of r e sp o n s iv ity o f each group tow ard th e fem ale stim u lu s i n a stan d ard sex t e s t s itu a tio n could be ev aluated* The change in s e x u a lity (freq u en cy o f "z ig -z a g s” ) fo r each of th e th re e groups o v er the t e n days of ex p erim en tatio n is p re se n te d g ra p h ic a lly in F ig u re 7* The t o t a l number of " z ig -z a g s ” toward th e fem ale stim ulus f o r each group was analyzed by means of a K rusk al-W allis a n a ly s is of variance* I t y ie ld e d an H « 6*1, *0j> > p > *02* Sub sequent a n a ly s is betw een In d iv id u a l groups showed th a t group R M gave more " z ig -z a g s” toward th e fem ale during th e d a lly sex te s ts th a n did th e group th a t re c e iv e d only th e sex t e s t durin g the te n days o f th e experim ent (Mann-Whitney T J = 10, p < *01). S im ila rly , group M M gave more resp o n ses toward the fem ale th a n group C (Mann- W hitney I T • 2l|.»0, p < .03)* T hus, In term s of th e over a l l d iffe re n c e s between groups comparing th e t o t a l number of " z ig -z a g s" summed over a l l te n days, both groups R M and M M were more resp o n siv e th a n group C* I t must be p o in te d out t h a t group M M was th e h ig h e st i n number of resp o n ses a t th e o u ts e t of th e experim ent alth o u g h th e o v e r a ll d iffe re n c e betw een groups on day one d id n o t Fig# 7 • —Change i n th e number of "z ig -z a g a " d ire c te d a t th e fem ale stim u lu s d u rin g sex t e s t a f o r a l l th re e groups over te n days* Each p o in t r e p re s e n ts th e average nz ig -z a g " freq u en cy p e r m inute change from th e i n i t i a l le v e l* M EAN RESPONSE CHANGE O V E R PAYS F R O M MITTAL LEVEL ( P A M 1) • R M 7 z 3 1 f * 7 8 9 s * & (INITIAL LEVEL) P A Y S 76 show th a t d iffe re n c e to be r e l i a b l e (K Tuskal-W allis H s 2 *8 , .3 0 p . 2 0) . A w ith in -g ro u p s a n a ly s is of each group, comparing day one frequency of 1 1 z ig -z a g s” w ith th e number on day te n , dem onstrated th a t only group R M showed a r e li a b l e In c rease from day one to day te n (W llcoxon T s 0 , p < .01). These an aly ses confirm th e im p ressio n g iv en by F ig u re ?• Group R M shows a r e la t i v e l y stead y in c re a se over th e course of th e experim ent i n th e le v e l of s e x u a lity d is played toward th e fem ale stim ulus* Group M M shows a le s s marked in c re a se up to day f i v e , but th e n re tu r n s to only a very s li g h tl y h ig h e r le v e l th a n i t m an ifested on day one. Group C shows no r e a l change o r tren d over the d u ra tio n o f the ex p erim en t. In summary, d a ily f i f t e e n m inute p re s e n ta tio n s of a r e a l male stim u lu s over a p e rio d o f te n days r e s u lte d i n an In c re a se In th e le v e l of ” z ig -z a g ” resp o n ses d ire c te d a t a fem ale stim u lu s d u rin g d a ily sex t e s t s . N eith er a d m in is tra tio n of th e model male nor th e no male trea tm en t re s u lte d in any m aintained change over th e te n days In th e le v e l o f ” z ig -z a g s” d u rin g sex t e s t s . C onclusions from th e Group E ff e c ts From the a n a ly s is o f th e d a ta tre a te d by groups, the e x p e c ta tio n th a t ag g ressiv e re sp o n se s would h a b itu a te w ith re p e a te d exposures to a norm ally a g g r e s s io n - e lic itin g stim u lu s I s w e ll su p p o rted . The f u r th e r p re d ic tio n t h a t "c h a rg e s” would wane b e fo re " b ite s " a ls o was supported* The fin d in g t h a t " o r ie n ta tio n s ” a lso waned was n o t ex pected* b u t th e f a c t t h a t th e y never dim inished to a v ery low le v e l even though th e decrem ent over days was s t a t i s t i c a l l y r e l i a b l e i s n o t s u rp ris in g i n l i g h t of th e f a c t th a t " o r ie n ta tio n s ” n e c e s s a rily precede b o th " b i te s ” and "c h a rg e s” and b o th o f th e s e resp o n ses showed very sh arp decrem ents* " O rie n ta tio n s ” are co n sid ered as re sp o n se s th a t n o t only precede " c h a rg e s," " b i t e s ," and " z ig - z a g s ," b u t a ls o as in d ic a to r s of v lg lla n c e » I t was p re d ic te d th a t d u rin g p r e s e n ta tio n s of th e m ale stim u lu s t h a t was b ein g h a b itu a te d th e amount of sex u a l b e h a v io r d ire c te d tow ard th a t stim u lu s would i n c re a se as th e a g g re ssiv e b eh av io rs began to decrease* Due to th e f a c t t h a t only to th e r e a l male stim u lu s was th e re any s ig n if ic a n t amount of sex u a l re sp o n siv e n e ss t h i s h y p o th e sis could be te s te d w ith in th a t group* This h y p o th e sis was supported i n th a t th e g re a t d e c lin e I n a g g re ss io n seen d u rin g day one was follow ed on day two by a s t r i k i n g upsurge I n sex u a l re sp o n se s to th e r e a l male* I t was r a th e r unexpected th a t t h i s e f f e c t would occur so soon i n th e ex p erim en t, b u t r e g a rd le s s o f how soon t h i s o ccu rren ce took p lace I t i s s tro n g evidence in su p p o rt of th e d ls ln h lb ltio n h y p o th e s is . F u rth e r su p p o rt 7 6 fop th e hypoth esized m utual in h ib ito r y d riv e I n te r a c tio n i s su p p lied by th e fin d in g th a t f o r group RM the le v e l of the seat d riv e as measured by th e number o f " z ig -z a g s ” to th e fem ale d u rin g sex t e s t s , In c rease d through th e d u ra tio n of the ex perim ent, whereas th e n o n -h a b itu ated c o n tro l group showed no change* In d iv id u a l S u b jects Hypotheses re g a rd in g th e occurrence o f d isp la c e ment a c t i v i t i e s stemming from th e e x te n sio n o f the d i s - in h lb itlo n h y p o th esis developed i n t h i s d is s e r ta tio n w ere c a s t in term s of p re d ic tin g displacem ent d u rin g a p e rio d of th e experim ent dem arcated on one hand by th e peak o f a g g ressiv e responses and th e peak of se x u a l behaviors d ire c te d tow ards th e same stim u lu s on the o th e r. In o th e r w ords, d isp lacem en t, p rim a rily in th e form of "fan n in g b o u ts ," should occur a f te r th e peak of a g g re ssiv e re s p o n s iv ity toward the r e a l male (o r model male in g ro u p M M ) but before th e peak o f sex u al r e s p o n s iv ity toward th a t stim ulus* Because th e peak occurrence o f sexual b ehaviors (" z ig -z a g s" ) during a g g r e s s io n -e lic itin g stim u lu s p re s e n ta tio n s was v a ria b le from s u b je c t to su b j e c t , th e p re d ic te d displacem ent could not reasonably b e expected to show as a group e ffe c t* Hence, t h i s e f f e c t m ust be sought w ith in each su b je c t tre a te d in d iv id u a lly * 79 O ther b eh av io rs o r fu n c tio n s p e c u lia r to in d iv id u a l sub je c ts w il l a lso be m entioned and commented upon in t h i s sectio n * R eal Male Group (RM) F ig u res 8 through 11 each c o n ta in g rap h ic p re s e n ta tio n s of th e d a ta f o r th e in d iv id u a l s u b je c ts in group R M * The d a ta p re se n te d f o r each s u b je c t are th e le v e l o f s e x u a lity (" z ig -z a g s" p e r m inute) as measured during th e m orning sex t e s t s (th e l in e h isto g ram in th e to p graph) , and f o r th e a fte rn o o n a g g r e s s io n - e lic itin g stim u lu s p re s e n ta tio n s , th e number o f "bites** p er m inute as averaged over su ccessiv e th re e m inute p e rio d s by days (th e p o in ts connected by a s o lid lin e in the top gr&ph)» th e number of "z ig -z a g s" p e r m inute as averaged over th re e m inute p e rio d s by days (th e p o in ts connected by a s o lid lin e in th e second g ra p h ), and th e amount of d is placem ent a c tiv ity p er m inute as averaged over su ccessiv e th re e m inute p e rio d s by days (th e d a r k - f ille d histo g ram a t th e bottom of the f i g u r e ) * The displacem ent a c ti v i t y graphed i s th e most fre q u e n tly o c c u rrin g one fo r th a t p a r tic u la r su b ject* F ig u re 8 p re s e n ts the d a ta fo r s u b je c ts 8ID and 13D-2. Number 8ID shows a sharp d ecrease in th e number of " b ite s " durin g the f i r s t day, w ith a more g rad u al change t h e r e a f t e r . The number o f "z ig -z a g s" durin g the 8Q F ig . 8 . —Graphic d a ta f o r f i s h number 8 ID and number 13D-2, group R M * F or form at e x p la n a tio n see te x t* / / r &/ 3 p v7~/ *1/ a / . / P £SP t > *< j es/ si u/ ure # 2 !D R H 9 0 70 C o S O ¥ y> 20 to C O * io 70 20 147 I1 7 10 067 0-3 } L I I I I i i i.i I n i i 1 n i i 1 > i . i I. i , . 1 . . >« I . . , , | t . . , 1 . . . . I Z li 1 I « i I > » ■ ■ 1 » n i I I I n f I n ■ . I I. 1 . . ttoiiMiii # /3D-2 RM vr § 40 0 N k ? _ * 3 o . x / r r * / 2 3 4 5 6 7 8 9 /tf DAYS 0 0 a g g r e s s io n - e lic itin g s tim u la tio n p e rio d s shows a r a th e r g ra d u a l in c re a s e to a peak re s p o n s iv ity on day s ix and th en a d e c re a se . On day two t h i s s u b je c t i n i t i a t e d h is maximum number of "fa n n in g bo u t s ^ T his f i s h , th e n , su p p o rts th e p re d ic tio n t h a t maximum d isp lacem en t w ill occur betw een th e peak of a g g re ssio n and th e peak o f se x u a l a c t i v i t y d ire c te d a t th e a g g r e s s io n - e lic itin g s tim u lu s , i n t h i s case th e r e a l m ale. The lin e h isto g ram in th e top graph in d ic a te s t h a t th e re i s a g ra d u al in c re a se in th e le v e l o f s e x u a lity d u rin g th e m orning sex t e s t s fo r th e e x te n t o f th e ex p erim en t. S u b je c t number 13D-2 shows le s s c le a r su p p o rt fo r th e h y p o th e s is . W hile th e maximum a g g re ssio n i s demon s tr a te d on day one d u rin g th e f i r s t th re e m inutes and the maximum se x u a l a c t i v i t y i s on day th r e e , th e maximum d a ily t o t a l o f disp lacem ent a c t i v i t y (" fa n n in g 1 1 ) does not < Both number o f bouts and th e d u ra tio n o f "fanning** were reco rd ed d u rin g th e ex p erim en t. A n aly sis i n term s of b o u ts o r d u ra tio n y ie ld e d e s s e n tia lly i d e n t i c a l r e s u l t s . The number o f b o u ts has been graphed th ro u g h o u t and used as th e m easure o f "fan n in g " a c t i v i t y . ^"Pushing" and "g lu in g " (d isp lace m e n t b e h a v io rs , a p p ro p ria te to n e a t-c o n s tru c tio n ) were fre q u e n tly engaged in by t h i s s u b je c t. However, t h e i r freq u en cy was le s s th a n "fan n in g b o u ts ," b u t showed th e same p a tte r n o f o ccurrence as "fan n in g b o u ts ." "Pushing" and "g lu in g " a re n o t f u r th e r an aly zed . 83 7 occur u n t i l day seven*1 However, I t i s w orth n o tin g th a t th e th re e m inute p e rio d r e f le c tin g th e g r e a te s t number of "fan n in g bouts" occurs during m inutes 7 * 8 and 9 on day one* Thus* w hile t h i s i s not evidence of stro n g support f o r th e h y p o th esis* i t i s su g g estiv e of an In c re a se o r peak of displacem ent a f t e r th e ra p id d e c lin e i n th e frequency of " b ite s " found a f te r th e f i r s t th re e m inutes of day one and b e fo re the maximum sex u al a c tiv ity * This f± 3h shows an in c re a s e i n se x u a l i t y in th e m orning sex t e s t s ov er the te n days o f th e experim ent* F ig u re 9 p re se n ts th e d a ta fo r s u b je c ts 9DP-5? and 12DP-5* S u b ject 9DP-5 a g a in does not com pletely su p p o rt the h y p o th esis re g a rd in g th e occurrence o f displacem ent a c t i v i t i e s b u t he does show a p a r t i a l e f fe c t* On day one th e le v e l of ag g ressio n tow ard the stim u lu s appears depressed although no d a ta from th e o r ig in a l re c o rd s nor from th e n o tes ta k e n during t h a t day in d ic a te any e x tra o rd in ary circum stances th a t m ight have r e s u lte d i n or caused such a depression* The In c re a se i n re s p o n s iv ity durin g day one r a th e r than th e decrease seen in o th e r 7 "Pushing" and "g lu in g " (d isp lacem en t behaviors* a p p ro p ria te to n e s t-c o n s tru c tio n ) were fre q u e n tly engaged i n by th is su b ject* However* t h e i r freq u en cy was le s s th an "fanning bouts*" b u t showed the same p a tte r n o f occurrence as "fan n in g b o u ts ." "pushing" end "g lu in g " are n o t f u r th e r analyzed* F ig , 9 . —Graphic d a ta f o r f i s h number 9DP-5> and number 12DP-5* group RM, For form at e x p la n a tio n see te x t* X9PP-5 4o 20 10 / I 3 9 £ £ 7 8 9 to fj V * -33 I ini[i iill iii 1 1 1 ni 1 j mli iuliiii li ii $0 £ 4o I \ 30 y > 10 A 6 * # C S T / C M 1 11 n 1 11111 ■■ 111 n 111 11 111 n 11 [ 111 11 1 in 1 1 / ^ 3 4 ? 6 7 8 PAY t t / i , , , i S r ^ r I W ? i ^ > )nl,rrr,A n r t ? T 1 ------- 1 ------- 1 ------- 1 -------- 1 — —i-------r C O \n s u b je c ts su g g ests th a t f e a r o f th e stim u lu s o r f r i g h t caused by th e in tro d u c tio n o f th e stim u lu s tu b e may have depressed the ag g ressiv e re sp o n se s* and perhaps* as th e f e a r waned th e s u b je c t became more aggressive* D uring th e l a t t e r th re e th ree-m in u te blocks th is s u b je c t engaged i n the only displacem ent t h a t i t showed d u rin g th e ex perim ent* On day two th e re appears an upsurge in th e frequency of sexual b eh a v io rs d ire c te d toward the r e a l male stim u lu s which reach es a peak on day six * Thus* th e maximum displacem ent does n o t ta k e p la c e a f te r the peak of aggression* b u t i t does o ccur befo re th e peak of s e x u a lity as d ire c te d toward the r e a l male* However* th e f a il u r e to com pletely confirm th e p re d ic tio n may have been due to confounding w ith a f e a r component th a t de p re sse d day one aggression* As w ith the p re v io u sly d is cussed su b je c ts in t h i s group* th e le v e l of s e x u a lity as measured by th e d a ily m orning sex t e s t s in c re a s e s over th e d u ra tio n of th e experim ent to a te rm in a l le v e l tw ice th a t found a t c r it e r io n (day one)* S u b ject number 12DP-5> does n o t give any evidence o f support f o r the displacem ent hypothesis* This s u b je c t shows a very h ig h le v e l of ag g ressio n on day one w hich i s follow ed by a ra p id decrease over th a t day* On day two th e maximum s e x u a lity and displacem ent a c tiv ity tow ards the r e a l male tak e place* This emergence o f displacem ent 87 m ust be in te r p r e te d in l i g h t of th e f a c t t h a t the b a rs on th e h isto g ram re p re s e n t only a t o t a l of two "fa n n in g bouts*" Even so* th e d isp lacem en t a c tiv i ty does n o t ta k e p la c e b efo re th e o n set o f th e peak of " z ig -z a g g in g ," b u t r a th e r d u rin g th e same p erio d * This s u b je c t a ls o shows th e ty p ic a l In c re a se in the number o f " z ig -z a g s ” d u rin g th e morning sex t e s t s over th e days o f the e x p e ri ment re a c h in g a f i n a l le v e l on day te n th a t i s tw ice th e o r ig in a l le v e l on day one* F ig u re 10 p re s e n ts th e d a ta f o r s u b je c ts 1 * .1 D and 71D* S u b je c t 1 j.1D shows a maximum " b itin g " freq u en cy on days one and two* a maximum "z ig -z a g g in g " tow ard th e same stim u lu s (th e r e a l m ale) on day six* w hich i s a ls o th e day o f f i r s t o ccu rren ce o f se x u a l b e h a v io r. T his s u b je c t d id not engage i n "fan n in g * ” but* as d e p ic te d on th e bottom graph* d id engage in an u n u su a lly la rg e amount o f "p ick in g " behavior,® C o n sid erin g th is "p ic k in g " b e h a v io r as a form o f displacem ent i t could r e p re s e n t d a ta b e a rin g on th e h y p o th esized tem p o ral order o f o ccurrence o f displacem ent betw een peaks of a g g re ssio n and sex* Q "P ick in g " i n th is c o n te x t re p re s e n ts a combina ti o n of th o se b e h a v io rs I d e n tif ie d and reco rd ed as "sand p ic k in g " and " m a te ria l te s tin g * " "Sand p ic k in g " and " m a te ria l te s tin g " are b o th a s so c ia te d w ith n e s t- o o n s tru c tlo n ; t h e i r occu rren ce d u rin g th e se x u a l phase may be reg ard ed as displacem ent* 88 Pig. 10.— Graphic data for fish number \\.XD and number 71D, group RM# For format explanation see text* # 4 / R M I n 11 Ii H I 111II i 1111 i 11 n II I 111 2 t -----1 — I — I ----r m Z /fr/M S V C M T f?fSPC/VS£S/h/W7£~ C Z 2 S O 30 1 0 i In ii 1 1u 11 ii n ) n ■■ 1 1 1 1 ■ It i ii In n lu -i-i-L 20 1 0 i . . . . r r r m r r r r i . . . . # 7 /0 m C O vO 90 However, I t d id n o t occur lie f ore the peak of se x u a l b e h a v io r d ire c te d toward the r e a l m ale; th e re fo re * I t m ust be conoluded th a t t h i s f i s h did n o t su p p o rt th e d is p la c e ment a c ti v i t y h y p o th esis* S u b je c t number 71D c e r ta in ly does n o t su p p o rt th e h y p o th e sis as he never engaged i n any d isp lacem en t a c t i v i t i e s * He re v e a ls a maximum number of " b ite s " on day one w ith a subsequent d ec rease and a peak o f " z ig zagging" on day f iv e to th e r e a l male stim u lu s* This s u b je c t shows an in c re a s e i n "z ig -z a g g in g " tow ard th e fem ale stim u lu s over th e te n days o f th e experim ent* F ig u re 11 p re s e n ts the d a ta f o r s u b je c ts 5>DP-£ and 7D* S u b je c t £DP-5 shows no d isp lacem en t a c t i v i t y d u rin g th e te n days o f ex p erim en tatio n * He d em o n strates ty p ic a l d ec rease o f " b itin g " from a peak on day one and a ls o a peak o f nz ig -z ag g in g n toward th e same stim u lu s on day fiv e * T his f is h a ls o shows the in c re a s e i n “ z ig zagging” over th e d u ra tio n of th e experim ent i n th e m orning sex te s ts * S u b je c t number 7D shows no d isp lacem en t a c tiv ity * He shows a g e n e ra l o v e r a ll d ec rease i n “b itin g " b eh av io r over the te n days o f e x p e rim e n ta tio n w ith th e r e a l male stim u lu s* The i n i t i a l le v e l o f " b itin g ," as r e f le c t e d i n th e f i r s t p o in t on th e g ra p h , fo r each of th e f i r s t th re e days shows a d ep ressio n * As w ith s u b je c t 9D P-S>> 91 Pig# 11#— Graphic data for fish number $DP-$ and number 7D» group RM# For format explanation see text* 10 1111111111111111111 il i ml m 111111 li 1 1 1 li m hi n I / 2 ? * 5 £ 7 8 9 10 D A Y I i l l m lltTili nH * n i l n n K m l n M #7-D 1 1 1 1 1 1 1 .1 1 1 ■ ! 1 1 1 1 ...i.1 1 1 1 .1 .1 1 1 1 .1 1 1 ,ii1 1 .. .it... 1X 1 DAYS 93 t h is d ep ression I s lik e ly due to an i n i t i a l fea r o f the stim ulus male in the tube th a t wanes a fte r the f i r s t couple of m inutes and thus r e s u lts in an in crea se in the number of "bites*" This f i s h shows l i t t l e "zig-zag*1 a c t iv it y d irected at the a g g r e s s io n -e lic itin g stim ulus* What "zig-zagging" a c t iv ity t h is f i s h displayed appeared during days seven and eig h t* Again, as w ith a l l the oth er su b jects in th is group, there i s an in crea se in the number of "zig-zags" d irected at the r e a l fem ale during th e morning sex t e s t s although th is su b ject shows the le a s t pronounced e f f e c t of th e f is h i n th is group* In summary, only one f i s h in group R M provides c le a r support fo r the ex ten sio n of th e d is in h lb ltlo n h yp oth esis which p red icts th a t the maximum frequency of displacem ent occurs between the peak o f aggression and the peak o f s e x u a lity as d irected towards the r e a l male during the a g g r e s s io n -e lic itin g stim ulus p resen tation s* Two a d d itio n a l f i s h give l e s s c le a r , b ut s u g g e s tiv e , support fo r th e hypothesis* Of the rem aining f i v e f is h on ly two showed displacem ent a c t iv i t ie s and n e ith e r showed the displacem ent at p eriod s during the experiment th a t would have been In terp reted as supporting th e exten s io n of the d ls in h lb ltio n h ypothesis* From the in d iv id u a l su b ject d ata graphs fo r th is group i t i s clea r th a t the decrease in "bites" depicted 9 k i n the p rev io u s s e c tio n o f t h i s c h a p te r in F ig u re If i s n o t an a r t i f a c t of av eraging hut r a th e r a fu n c tio n shared by a l l th e members o f th e group* S im ila rly , th e group d a ta graph t h a t shows th e In c re a s e in "z ig -z a g s" over days in th e morning sex t e s t s (F ig u re 9) i s a lso a fu n c ti o n common to every member o f group R M * Model Male Group (M M ) F ig u re s 12 through lj? each c o n ta in g rap h ic p re s e n ta tio n s o f th e d a ta f o r th e in d iv id u a l s u b je c ts i n group M M * The form at of th e se fig u re s i s id e n tic a l to t h a t f o r group RM * F ig u re 12 p re se n ts th e d a ta fo r s u b je c ts number 10D and 3D* Number 10D i s th e only f i s h i n group M M th a t engaged i n displacem ent b e h a v io rs; i n t h i s case i t was "p ic k s" t h a t emerged* The i n i t i a l number of " b ite s " d ire c te d a t th e model i s maximal on day one* T his peak i s r a th e r low, re ach in g only a r a te of te n " b ite s " p er m inute d u rin g m inutes k* $ and 6* The number o f " z ig zags" d ire c te d a t th e model i s g r e a te s t on day s i x , d is re g a rd in g th e "z ig -z a g s" on day e lev e n which occurred d ire c te d toward a r e a l male d u rin g th e r e a l male p o s t- te s t* The maximum number o f "p ic k s" (th e displacem ent a c ti v i ty ) occurs on day th re e * Thus, th e only f i s h i n group M M to engage in displacem ent a c t i v i t i e s does so in a manner t h a t su p p o rts th e p re d ic tio n of th e tem poral 95 Pig* 12*-—Graphic data for fish number 10D and number 3D, group MM* For format explanation see text* v O O ' S A V O 0 / 6 8 6 J i k £ Z / m iiiMiiirTTim im n n i M i m iin iim iiinnr m u m iiinfiunim iniq 1 0/ % 0£ I O j * h i < * I (IK M J 05 09 T Y u i i i i i , T 1 * ^ **$ v T n ^ i i s a v o 0 ! G 2 L 3 ^ ^ 7 ^ Z / U * § ^Tjn^ 1 1 1 1 IjlJ* I1 0! c?z§ 1 0 £ § t 0* * 97 r e la tio n s h ip s of th e o ccurrence of th e peak freq u en cy o f a g g re ss io n , d isp lacem en t, and sex u a l a c t i v i t i e s d u rin g th e a g g r e s s io n - e lic itin g stim u lu s p re s e n ta tio n s * This f i s h shows a s lig h t d e c lin e i n th e number o f " z ig -z a g s" toward a fem ale d u rin g th e sex te s ts * The p o s t - t e s t p r e s e n ta tio n o f a r e a l male on day e le v e n shows a h i gh le v e l of r e s p o n s iv ity ( " b ite s ” ) toward th e stim ulus* T his f i s h a ls o re a c te d to th e r e a l male w ith a number of " z ig -z a g s ” g re a te r th a n th e number d ire c te d toward th e model on any day o f th e experim ent* S u b je c t number 3D shows a peak o f " b itin g " on day one follow ed by a t y p ic a l d ecrease on th e days fo llo w in g * The peak o f "z ig -z a g g in g " fo llo w s on day six* The le v e l o f " z ig -z a g s ” during sex t e s t shows a s l i g h t in c re a s e over the d u ra tio n of th e experim ent* The p o s t - t e s t p re s e n ta tio n of the r e a l male r e s u lte d i n a v ery h ig h le v e l of a g g re ssio n ( " b ite s " ) toward t h a t stim ulus* F ig u re 13 p re s e n ts th e d a ta f o r s u b je c ts f> D and 11D-2. Humber $D i s unique i n t h a t , d u rin g the f i r s t fo u r d ay s, th e re are a la rg e number o f " th r e a ts " and d u rin g th e s e fo u r days th e re i s an in c re a s e i n th e number o f " b ite s ” d ire c te d a t th e model* However, a f te r day th re e th e re i s a d e c re a se in th e freq u en cy of " th r e a ts " and by th e end of day fo u r " b ite s " has reach ed i t s peak and th e y , to o , b eg in to d ecrease over th e rem ain in g days 98 m x m h a y . ? f s * x3*~-G raphic d a* a fo r f i s h number 5d and number l l D - 2 , group mm. Por form at e x p la n a tio n s ^ T t e x t . t u r e a j s / m in . r e s p o n s e s / MIH- 70 • G O 50 A O 50 20 10 i f SO MM <? (r e a l ) l u i i l u - u - l i m l i i i i l m i l i i i i l i i i i l i i i i l i m l i n i l i m 10 f I I II il l II t i l l I i l l I l l l l l I l l I I I ll I H I I I I I I I H 1 I I I II if 1 11 1 / 2 ? * 5 € 7 g 9 / 0 / / DAYS #HD‘ l m n n l n . i l i m l m i h i n l n i i l i n i l i n . l n M l m i I n n DAYS vO vO ioo o f th a ex p erim en t. This f i s h d id n o t respond to th e model w ith se x u a l behavior* The le v e l of ” z ig -z a g s ” d ire c te d a t th e fem ale d u rin g th e sex t e s t s shows a v ery sh arp in c re a s e over th e d u ra tio n o f th e ex p erim en t• The p o s t - t e s t p re s e n ta tio n o f th e r e a l m ale produced a h ig h le v e l of ag gression* S u b je c t number 1XD-2 shows a peak of ”b i t e s ” a t a low le v e l on day one and very few more on subsequent days* T his f is h d id n o t ” z ig -z a g ” toward th e model* D uring th e sex t e s t s th e le v e l of ” z ig -z ag g in g ” f lu c tu a te d b u t began and term in ated a t e s s e n t ia l ly th e same le v e l* The p o s t - t e s t w ith th e r e a l male stim u lu s e l i c i t e d a ty p ic a l h ig h le v e l of aggression* F ig u re 1^ p re s e n ts th e d a ta f o r s u b je c t number 11D and 63D. Number 11D shows the low est le v e l o f a g g re ssio n of th e sample of f i s h used i n t h i s s tu d y . Peak ”b itin g ” i s found on day one, th e le v e l never exceeding more th a n f iv e ”b i t e s ” p er m inute* T h e re a fte r, th e frequency o f ”b l t e s ” d e c re a se s to a n ear zero le v e l f o r th e rem ainder of th e experim ent* The number o f ” z ig -z a g s” toward th e fem ale stim u lu s d u rin g sex t e s t s shows a s l i g h t d ecrease over th e d u ra tio n of th e experim ent from an i n i t i a l le v e l which i s th e lo w est o f any f i s h i n group M M * P re s e n ta tio n of the r e a l male in th e p o s t - t e s t r e s u lte d in a m oderate le v e l of aggression* 101 Pig# 14.— Graphic data for fiah number 1XD and number 63D, group MM# For format explanation see text# #///> m 40 10 M.i.Mtir/SirTrTrtttiTrttttmrmtrYTnTTTtii.... / 2 3 4 5 6 7 8 S fo // DAYS #0-5-0 M M Z0 " I m i l l n In nil in 11 ii il ii i ill im I n n li.i. Ii. ii In ii 103 Number 63D shows a p e c u lia r ity I n th a t th e le v e l of a g g re ssio n shows an In c re a s e w ith in each day r a th e r th an th e more c h a r a c te r is tic decrease* The number o f " b ite s " p e r day does show a d ecrease over days* The number o f " z ig -z a g s" p e r m inute d u rin g the m orning sex t e s t s shows a s lig h t in c re a s e over th e te n days o f th e experim ent* P re s e n ta tio n of th e r e a l male on th e e le v e n p o s t - t e s t r e s u lte d i n a h ig h number o f "bites® d u rin g a l l m inutes excep t m inutes sev en , e ig h t and nine* The sudden c e s s a tio n and reap p earan ce o f " b itin g " I s u n ex p lain ab le from th e data* The re c o rd f o r t h i s s e s s io n and th e accompanying n o tes g iv e no in d ic a tio n of any abnormal ev en ts t h a t would e x p la in such a sudden change* T his f i s h did n o t "z ig -z a g " toward th e model d u rin g th e ex perim ent nor show any d isp lacem en t a c tiv ity * F ig u re 1^ p re s e n ts the d a ta f o r s u b je c ts 9^ an^ Lp* F ish number 9® shows a d e c lin e over th e d u ra tio n of the experim ent in the number o f " b ite s " from a peak on day one* T his f i s h does n o t show any change o th e r th an d a ily f lu c tu a tio n s i n th e le v e l o f s e x u a lity d ire c te d a t th e fem ale stim u lu s d u rin g th e experim ent* This f i s h d id n o t " z ig -z a g " toward th e model n o r show any displacem ent* S u b je c t I p shows a p a tte r n s im ila r to 9D ex cep t t h a t , w hile th e peak number o f " b ite s " does occur to th e model on day one, th e d im in u tio n of " b itin g " does not Pig* 15«—Graphic d a ta fo r f i s h number Q D and number 1 4 D* group T .- T M * Por form at e x p la n a tio n see te x t* 1A O H SAVO f l O l £ 9 L 9 $ i t £ Z I TTTTT fii< iill 1 1 1 1 rr1 1 m v 1 1 1 1 1 1 1 1 1 1 1 ii1 1 it 0/ OZ oc Ob 0$ 03 J OL Ntu/ S 3 $ N O d S 3 y I I Q I G Q L B Z + ST! 111 H 1 1 111 M IIJI111IU 1^1 1II11111111111| H TT' I rmfTT T11 ‘ i» ,A ' Y (7 V3 $ ! t ww 0~l,& o / OZ o i Ob o$ 03 'NW/s'3SN0d$3& 106 prooeed as ra p id ly * Hot u n t i l day e ig h t I s th e re a r e - d u o tlo n to n ear zero le v e ls* The le v e l o f s e x u a lity shown d u rin g th e sex t e s t s shows some f lu c tu a tio n s b u t ends a t approxim ately th e same le v e l as I t s t a r t s * T his f i s h n e ith e r "zig -zag g ed " tow ard th e model nor showed any d isp lacem en t a c tiv ity * The a g g re ssiv e resp o n se to th e p o s t - t e s t p re s e n ta tio n of th e r e a l male was h ig h e r th a n by any o th e r member o f t h i s group* In summary, only one f i s h i n group M M showed any disp lacem en t a c ti v i t y w hatsoever* That f i s h provided su p p o rt f o r th e e x te n s io n o f th e d is in h ib ltio n h y p o th e sis t h a t i s , th e maximum frequency of disp lacem en t b eh a v io r oocurred betw een th e peaks of a g g re ssio n and th e pealt o f s e x u a lity i n resp o n se to th e model male* No o th e r f i s h responded w ith d isp lacem en t and o nly one o th e r showed any se x u a l re sp o n se s to th e model* Prom th e in d iv id u a l s u b je c t graphs f o r t h i s group i t i s c le a r t h a t th e decrem ent i n a g g re ssiv e re sp o n se s over days shown i n th e g ra p h ic p re s e n ta tio n (F ig u re 10 i n th e p reced in g s e c tio n i s n o t an a r t i f a c t of averaging* The d ecrease from day to day i s c h a r a c t e r is t ic o f seven of th e e ig h t f i s h i n t h i s group* Only the f i s h t h a t showed a la rg e number of " th r e a ts " to th e model d u rin g th e f i r s t th re e or fo u r days may be co n sid ered to be a ty p ic a l of th e group fu n c tio n (number £D)* M l I n th e p rev io u s s e c tio n ("Group E f f e c ts 1 1 ) o f th is c h a p te r th e fin d in g th a t th e re was no d iffe re n c e betw een th e day e lev e n re a l-m a le p o s t - t e s t resp o n se le v e l o f t h i s group and th e f i r s t day of stim u lu s p r e s e n ta tio n of th e r e a l male group was in te r p r e te d as su p p o rtin g a stim u lu s s p e c if ic it y o f th e decrem ent found w ith group M M to th e model male over th e e x te n t of th e experim ent* The h ig h le v e l of resp o n se to th e r e a l m ale i s found in a l l e ig h t of th e s u b je c ts i n t h i s group* I n a l l e ig h t f i s h th e le v e l of resp o n se to th e r e a l m ale on day elev e n exceeds th e resp o n se le v e l to th e model d u rin g any of th e f i r s t te n days* Control Group F ig u re s 16 through 19 each c o n ta in g rap h ic p re s e n ta tio n s of th e in d iv id u a l s u b je c ts in th e c o n tro l group (C)* The form at of th e se fig u r e s i s th e same as th e p rev io u s two groups d iscu ssed except t h a t th e y are much a b b re v ia te d in th a t only two m easures are re c o rd e d : th e le v e l o f s e x u a lity (" a ig -z a g s " ) as seen In d a ily morning sex t e s t s and th e re s p o n s iv ity to th e r e a l male i n th e day te n a fte rn o o n p o s t- te s t* F ig u re 16 p re s e n ts th e d a ta f o r s u b je c ts number 3D and number 2D-2* Number 3D shows a s l i g h t d ecrease i n sex t e s t resp o n se le v e l over th e te n days of th e experim ent* The p o s t - t e s t p re s e n ta tio n of th e r e a l male 108 Sis£ ^.--Graphic data for fish number 3D and number 2D-2, group C* For format explanation see text* 4 4 4 4 4 4 109 0 ( T > 0 0 C " v o U N '4- r < \ 01 o o o o ^ ^T\ K\ C\l -N iu / $3SM 0dS?X Q N \ < H “ J __L < 5» <r» 0 0 f- VO Vr\ £ 5 Q ■ ’ i' <M W W /S3S/V O c/S3& DAYS X10 r e s u lte d In a m oderate le v e l o f a g g re ssio n (" b ite s* ')* S u b je c t number 2D-2 shows an I n c re a s e , q u ite la r g e , i n the number of "z ig -z a g s" m a n ife st d u rin g the d a lly sex te s ta * The p o s t - t e s t w ith th e r e a l m ale r e s u lte d i n about th e same le v e l of a g g re ssio n as found w ith number 3D* F ig u re 17 p re s e n ts th e d a ta f o r s u b je c ts number 2D and !L D P-5>• Number 2D shows a s l i g h t in c re a s e I n th e le v e l o f s e x u a lity ov er th e te n days of th e experim ent* The p o s t - t e s t w ith th e r e a l male r e s u lte d in a v ery low frequency of ''b ite s * d ire c te d a t th e male in th e tube* T his number of " b ite s * i s th e few est d ire c te d a t th e r e a l male by any f i s h on th e p o s t- te s t* S u b je c t number 1DF-5 shows a s l i g h t d ecrease in th e le v e l of s e x u a lity over th e d u ra tio n o f th e e x p e ri ment* The r e a l male p o s t - t e s t p re s e n ta tio n e l i c i t e d s tro n g aggression* F ig u re 18 p re s e n ts th e d a ta f o r s u b je c ts ll(£-2 and 13DP-5>* Number llt-D-2 shows a marked d e c lin e i n th e number of " z ig -z a g s" d ire c te d a t th e fem ale stim u lu s d u rin g the te n d a ily sex te s ts * The r e a l male p re s e n ta t io n durin g th e p o s t - t e s t on day te n r e s u lte d i n a la rg e number of " b i t e s •" In s te a d o f s t a r t i n g a t a h ig h le v e l d u rin g th e f i r s t th re e m inutes and d e c re a sin g th e r e a f te r * th ey s ta r te d low and in c re a s e d over th e f i r s t tw elve XIX Fig* X7*— Graphic data for fish number 2D and number 1DP-5* group C. For format explanation see text. ^ 20 10 k 30 zo ll 111111111111 > 11111111111111111111111111111111111 113 rm**™ fife l8*"“GraPhic data for fish number lltfD-2 and number 13DP-5, group c. For format explanation see text* # W -z P / ? £ S P O t f S £ s/ / ? W T £ r #13 DP-5 m i Ii ii 111 m i n i 111 m i l n 111 m l n i ill m i n i i 9 JO DAYS H & ll£ m inutes* S u b ject number 13DP-5 shows no change in th e le v e l of s e x u a lity over th e ex p erim en t. P re s e n ta tio n o f th e r e a l male in th e p o s t - t e s t produced a la rg e number of " b i t e s ” d u rin g th e f i r s t th re e m inutes and a d ecrease th e re a fte r* F igure 19 p re s e n ts th e d a ta fo r s u b je c ts number ID and 12D* Number ID shows a very v a r ia b le le v e l of s e x u a lity from day to day. Comparing th e f i r s t day w ith the l a s t th e re i s a s l i g h t In c re a se in th e le v e l of s e x u a lity a f te r te n d ay s. The p o s t - te s t p re s e n ta tio n of th e r e a l male in th e tube r e s u lte d in an e x tra o rd in a ri ly higjh le v e l of " b ite s " d u rin g th e f i r s t s ix m inutes and th en a ra p id d e c lin e th e re a fte r* S u b ject number 12D shows a g radual b u t c o n s is te n t in c re a se in th e number of "z ig -z a g s" e l i c i t e d d u rin g th e d a lly sex t e s t s . The r e a l m ale, p re se n te d during th e p o s t - t e s t , e l i c i t e d a r e l a ti v e l y h ig h le v e l of a g g re ssio n which showed some In c re a se over th e f i f t e e n m inutes of th e t e s t r a th e r th a n th e more u s u a l decrease* In summary, t h i s group was p re se n te d w ith th e te n d a ily morning s e x - te s t. The purpose of t h i s procedure was tw o fo ld ; f i r s t , i t prov id ed a b a s e lin e fo r th e hypothesized changes in s e x u a lity th a t h a b itu a tio n of ag g ressio n might c re a te in the o th e r groups, and second, Pig* 19*— G raphic d a ta fo r f i s h number ID and number 12D, group C* For form at e x p la n a tio n see t e x t . M r s RESPONSE'S/ MIN. ^ « > w o o O o o o o o oo < P V o v * l § vn O D C X ) s R E S P O N S E S /n /M ro V - ^ - * s n « r > o Q > o o o c > O O T 1 ------- 1 - - - - - - 1 ------- 1 ------- 1 ------- 1 ------- 1 -------1 -------1 ------- 1 ------- 1 ------- 1 - 1 LXX QZ/# 118 a l l members o f t h i s c o n tr o l group were ad m in iste red a p o s t - t e s t p re s e n ta tio n o f th e r e a l male stim u lu s* Both group a n a ly s is and s u b je c t by s u b je c t a n a ly s is shoved th a t th e re was no c o n s is te n t in c re a s e o r d ec re a se in th e le v e l of s e x u a lity over th e d u ra tio n of th e experim ent* Comparing the le v e l of a g g re ssio n e l i c i t e d i n t h i s p o s t t e s t w ith th a t found on day one w ith group R M in d ic a te d th a t th e te n days th a t passed d u rin g th e experim ent were n o t, I n and of th e m se lv e s, s u f f i c ie n t to b rin g any d ecrease in th e le v e l o f agg ressio n * A ll e ig h t o f th e f i s h i n t h i s group (C) responded a g g re s s iv e ly tow ard th e r e a l male in th e p o s t- te s t* CHAPTER VI DISCUSSION The purpose of t h i s re se a rc h was twofold* F i r s t , an attem p t was made to extend some e a r l ie r fin d in g s w ith b ird s (H Inde, 1 9 5 ; 1961; M elzaok, 1961) In d ic a tin g th a t re p e a te d ly e l i c i t e d fix e d a c tio n p a tte r n s (FAPs) waned o r h ab itu ated * These r e s u l t s extend to more com plicated b eh av io rs such as FAPs and groups of FAPs subsumed under the heading of a " c la s s of b eh av io rs" (such a s "sex" or " a g g re s s io n " ), a fin d in g t h a t Is r a th e r common w ith re f le x iv e re sp o n se s (see H a r r is , 19^3; Thompson and S pencer, I 966) • S ev eral c r i t e r i a must be a p p lied to any observed decrem ent i n re s p o n s iv lty to an unconditioned stim u lu s In o rd er to c le a r ly d if f e r e n tia te i t from fa tig u e and sensory a d a p ta tio n b e fo re the phenomenon may ju s t if ia b ly be la b e le d h a b itu a tio n . Secondly, a re c e n t th eo ry of d riv e in te r a c tio n , the d l s - I n h lb itio n h y p o th esis of van I s r a e l and Bol (195>8) and S ev en ster ( 1961) , s tr e s s e s re c ip ro c a l in h ib itio n of d riv e s whloh ( i f one c o n sid e rs th e p ro cess u n d erly in g h a b itu a tio n to be one o f in te r n a l in h ib itio n of d riv e ) would p re d ic t s p e c if ic changes in behavior I n the p ro c e ss 119 120 o f h a b itu a tin g a stim u lu s. F u rth e r, th e p re se n t au th o r has sought to extend and m odify th e d is ln h lb ltlo n h y p o th esis t o p r e d ic t f u r th e r changes in b eh a v io r; th a t i s , th e occurrence o f displacem ent a c t i v i t i e s during ag g ressiv e defense of t e r r i t o r y in th e th re e -a p in e d s tic k le b a c k . The r e s u l ts o f t h i s stu d y w i l l be d iscu ssed under th re e h ead in g s: h a b itu a tio n of fix e d a c tio n p a t te r n s , th e d is ln h lb ltlo n h y p o th e s is , and th e e x te n sio n of the d is ln h lb ltlo n h y p o th esis* H a b itu a tio n of F ixed A ction p a tte rn s There i s no q u e stio n th a t b oth th e groups p re sen ted w ith a g g r e s s io n - e lic itin g s tim u li ( r e a l male or model) f o r f i f t e e n m inutes over te n days shows a d ecre ment i n a g g re ssio n over th o se te n d ay s. T his decrem ent may be term ed nh a b itu a tio n ,t In s o fa r as i n b o th cases the decrement i s p erp etu ated over th e tem poral g u lf betw een d a lly p re s e n ta tio n s , th u s d isp la y in g th e tem poral p e rs is te n c e c r it e r io n o f h a b itu a tio n . Temporal p e r s i s t ence would n o t be seen i f th e decrem ent were due to e ith e r fa tig u e or sensory a d a p ta tio n . F urtherm ore, th e group p re se n te d w ith th e model shows a stim u lu s s p e c i f i c i t y of th e h a b itu a tio n as shown by th e day e le v e n p o s t - t e s t p re s e n ta tio n of th e r e a l m ale w hich e l i c i t e d a h igh le v e l of a g g re ssio n . T his stim u lu s s p e c if ic it y 121 t e s t was n o t ap p lied to th e group th a t was h a b itu a te d to th e r e a l male because the p re -a d a p ta tio n procedure w ith th e wooden "blank model" could not be accom plished between th e day te n r e a l m ale p re s e n ta tio n and th e day e le v e n p o s t- te s t* The a g g r e s s io n - e lic itin g s tim u la tio n p erio d of f i f t e e n m inutes d u ra tio n was s e le c te d r a th e r a r b i t r a r i l y as a s tim u la tio n p erio d somewhat lo n g er th a n th e p e rio d used fo r an "a g g re ssio n te s t" * - i n th e e th o lo g ic a l l i t e r a t u r e (fiv e m inutes) and s h o rte r th a n th e tw enty to t h i r t y m inutes p re v io u sly used i n a p re lim in a ry stu d y of h a b itu a tio n by wyers (p e rso n a l com m unication), and by van I s r a e l in h is study of th e f a c to r s c o n tr o llin g p a re n ta l b ehavior (195>3)* The f iv e m inute a g g re ssio n t e s t does n o t r e s u l t i n h a b itu a tio n w ith re p e a te d d a lly p re se n ta tio n s * F ig u re 1 (C hapter I I ) was c o n stru c te d and schem atized from a la rg e body o f d a ta from th e la b o ra to r ie s a t the U n iv e rsity of Leiden by use o f r e peated measurements over days* Repeated t e s t s over days have been used i n o th e r s tu d ie s (e*g*„ van l e r s e l , 1953) and no r e p o rt has ever been made in d ic a tin g a d ecrease i n 1An "a g g re ssio n t e s t " i s ty p ic a lly a f iv e m inute p re s e n ta tio n of a r e a l male i n a c le a r tu b e p laced in a stan d ard p o sitio n * I t i s th e same as a "sex t e s t " w ith th e s u b s titu tio n of a r e a l male f o r a fem ale stim ulus* (See S e v e n ste r, I 96I , page 16, f o r a d e ta ile d d e s c rip tio n of the "a g g re ssio n te s t* " ) 122 re sp o n siv lty * However, i t should be m entioned th a t w hile e th o lo g is ts observe and re c o rd a l l b eh a v io rs th a t they can reo o g n ize and d e f in e * th e y tend to p re se n t t h e ir r e s u l t s i n term s of a dominant behavior* one u s u a lly most fre q u e n tly o r most r e lia b l y observed* In th e case of th ese f iv e m inute a g g ressio n t e s t s the r e s u l t s a re p resen ted in term s of b ite s * I t was shown i n th e p re se n t study th a t b ite s h a b itu a te d a t a much slow er r a te th a n th e most In te n se response* charges* which h a b itu a te d r a th e r quickly* While b i t e s do n o t h a b itu a te in stan d ard f iv e m inute ag g ressio n te s ts * th e re may be some decrem ent i n charges which would in d ic a te t h a t one o f the component beh av io rs in ag g ressio n i s s u s c e p tib le to h a b itu a tio n w ith sh o rt s tim u la tio n p erio d s* T his q u e stio n aw aits an answer i n f u tu r e research * In term s of the two m easures of a g g re ssio n used i n th is study* charges and b ite s* i t was found th a t both o f th ese components of th e beh av io r c la ss h a b itu a te d * and fu rth e r* th e y h a b itu a te d a t d i f f e r e n t i a l r a t e s ; th a t is* th e more ag g ressiv e response* charges* h a b itu a te d f i r s t follow ed by b ite s * These r e s u l ts su p p o rt the fin d in g s o f Hinde ( 195> l 4 .a, b ; 1961)* Melzaek (1961)* and s z le p ( I 96I 4 .)* where a PAP was found to h a b itu a te w ith re p e a te d e l i c it a ti o n * The stim u lu s s p e c if ic ity I n group M M and th e tem poral p e rs is te n c e found w ith b o th groups shows a b asic s im ila r ity between th e h a b itu a tio n o f FAPs and sim p ler r e f le x iv e responses* F urther* t h i s study extends th e fin d in g o f h a b itu a tio n t o a p h y le tic le v e l where v ery l i t t l e work has been done w ith t h i s phenomenon (T horpe, 1963)* A re c e n t stu d y by B aenninger ( 1966) p u rp o rts to dem onstrate h a b itu a tio n of ag g ressio n w ith Beta splanding (th e Siamese F ig h tin g Fish}* b u t th e method used by th a t author i s not s u f f ic ie n t to re a c h a c le a r c o n c lu sio n th a t what was dem onstrated was h a b itu a tio n r a th e r th an sim ple f r i g h t of the stim u lu s (see Appendix A f o r more d e ta ile d d isc u ssio n of t h i s study) * Ho d em o n stratio n of tem poral p e rs is te n c e was provided and no attem pt to show stim u lu s s p e c if ic ity was included* M elzaek, e t a l * ( I 96I) and Melzaek ( I 961) found th a t th e f r ig h t resp o n ses of ch ic k s waned w ith re p e a te d e l i c i t a t i o n from a s ilh o u e tte of a M haw k-liken shape flow n overhead* However, he found th a t th ey n e v e r, w ith in th e con fin es of h is experim ent, stopped o rie n tin g toward the stim ulus* They rem ained v i g i l a n t, b u t no lo n g e r responded w ith o v e rt fe a r* In th e case of the p re s e n t 3tudy an attem p t was made to m easure o rie n tin g resp o n ses In th e hope of p ro v id in g a comparable measure In t h i s s itu a tio n * I t was found th a t th e re was a d ecrease in the number o f o rie n ta tio n s throughout the experim ent* How e v e r, as was p o in ted out i n th e preceding chapter* t h i s decrem ent may have r e f le c te d only a change i n th e ag g res siv e resp o n ses which* by th e d e f in itio n s of th o se r e sponses used i n t h i s study* were n e c e s s a rily preceded by an O r i e n ta ti o n . ’ 1 The ag g ressiv e resp o n ses were reduced to very low le v e ls by re p e a te d e l i c i t a t i o n th ro u g i stim u lu s p re s e n ta tio n s w hereas the o rie n ta tio n s never reaohed a v ery low l e v e l . I t would be pure s p e c u la tio n to suggest t h a t they never would have waned com pletely* b u t i t does seem th a t they a re much more r e s i s t a n t to h a b itu a tio n th a n o v e rt ag g ressiv e re sp o n se s. To t h i s ex ten t* then* th e p e rs is te n c e of v ig ila n c e resp o n ses may be sa id to be s im ila r in th e case of o rie n ta tio n s toward a c o n sp e c iflc a tta c k e r when compered w ith the d if f e r e n t- sp eo ies p re d a to r o rie n ta tio n response i n M elzack's ( I96I) stu d y . Szlep (19&J.)* stu d y in g prey c a tc h in g resp o n ses i n th e sp id e r to v ib ra to ry s tim u la tio n o f th e webb* found th a t o rie n ta tio n s ( np e rc e p tio n s of th e stim u lu s1 1 ) would wane co m p letely . O rie n ta tio n s were th e f i n a l resp o n se to wane in th e complex o f responses d ire c te d a t th e v ib ra to ry stim u lu s designed to Im ita te prey c a u g it in the webb. In t h i s study* i n c o n tra s t to Hinde ( l g ^ a , b; 1961) , Melzaek e t a l . ( I 96I ) , Melzaek {I96I ) and th e p re se n t experim ent* th e ^ h a b itu a te d ” stim ulus was the prey and th e organism s tu d ie d , the p re d a to r. One marked d iffe re n c e between th e r e s u l t s of th is stu dy of h a b itu a tio n o f a PAP and th e u s u a l fin d in g s w ith r e fle x iv e response h a b itu a tio n was th a t th e stim u lu s e l i c i t i n g th e g r e a te s t ag g ressio n h a b itu a te d a t a f a s t e r r a te th a n th e le s s adequate stim ulus* The ty p ic a l f in d in g w ith re fle x iv e resp o n ses i s th a t th e le s s in te n s e s tim u li h a b itu a te more ra p id ly * A ggressive resp o n ses to the r e a l male d eclin ed a t a much s te e p e r g ra d ie n t and f i n a l l y reached alm ost th e same te rm in a l le v e l as the resp o n ses to th e model* This would su g g est th a t th e re m ight be a t le a s t one m ajor d iffe re n c e in th e h a b ltu a to ry p ro cess between FAP and r e f l e x h a b itu a tio n * A second d iffe re n c e between h a b itu a tio n of FAPs and sim ple re f le x e s i s t h a t FAPs ten d to ta k e much lo n g er to h a b itu a te * This may be m isleading* however* and may n o t r e a l l y r e f l e c t a b a sic d iffe re n c e betw een r e f le x and FAP h a b itu a tio n * When th e h a b itu a tio n of a FAP i s d is cussed* as in the p re s e n t context* one fin d s th a t components of the beh av io r in q u e stio n h a b itu a te in an ordered r e la tio n s h ip ; th a t is * one component of a g g re ssio n (e*g** charg es) w ill wane b e fo re th e n ex t (e,g** b ite s ) and so on* T his kind o f ordered r e la tio n s h ip was a prom inent fin d in g in th e d e ta ile d work w ith th e s p id e r re p o rte d by S zlep ( I 96J+ ) • I f I t i s assumed th a t each component i n t h i s ordered r e la tio n s h ip h a b itu a te s in a sequence and i f th e speed and course o f h a b itu a tio n of each component i s stu d ie d s e p a ra te ly i t m ight w e ll be th a t th e components o f a FAP and r e fle x e s would show s im ila r courses of response decrement* Such a r e s u l t would su g g est a common mechanism a t some fundam ental le v e l, perhaps m o le c u la r, sub summing a l l h a b itu a tio n phenomena* Support f o r such an I n te r p r e ta tio n and p o s tu la tio n aw aits s tu d ie s designed to f i r s t f r a c tio n a te , through c a re fu l o b serv a tio n of th e t o t a l b e h a v io r, component resp o n ses o f complexes such as fix e d a c tio n p a tte r n s , and th en re co rd th e waning o f each o f th e se components* A f i n a l th e o r e tic a l problem d eserv es b r ie f m ention and d is c u ss io n b e fo re proceeding to th e d isc u ssio n o f th e im p lic a tio n s of h a b itu a tio n of a g g re ssio n fo r th e d is - in h ib itio n hypothesis* This i s the q u e stio n of th e fu n c tio n a l locus of th e mechanism(s) su b serv in g h a b itu a tio n of fix e d a c tio n p a tte rn s * Is t h i s a re c e p to r p ro c e s s , a c e n tr a l p ro c ess o r a fu n c tio n re s id in g i n the ex ecu tiv e motor system ? I f th e resp o n se decrem ent ob served in t h i s study were due to a change i n th e a b i l i t y of th e f i s h to p e rc e iv e th e stim u lu s, th e f i s h would g ra d u a lly sto p resp o n d in g to th e h a b itu a tin g stim ulus* However, th is i s n o t a l l th a t happened in the course of th e h a b itu a tio n * The m ale n o t only decreased h is 127 a g g re ssiv e re s p o n s iv ity b u t shoved a q u a lita tiv e change In h is response to th a t stim u lu s, t h a t i s , sex u al be h a v io r emerged* There i s no re c e p to r p ro cess known th a t would e x p la in such a q u a lita tiv e change* S im ila rly , no m otor mechanism can e x p la in a q u a lita tiv e change in response such as t h a t found i n th e p re se n t study* Al though sim ple c e s s a tio n of response m ight be explained by in h ib itio n on th e e f f e r e n t o u tp u t sid e of th e c e n tr a l nervous system , th e emergence of a new response would argue th a t m otor in h ib itio n alone cannot e x p la in th e e n t ir e process* By e lim in a tio n t h i s leav e s the mechanism r e s id in g i n a c e n tr a l process* M arler ( I 96I ) , i n a paper concerned w ith th e s p e c if ic ity of re sp o n siv e n ess in in s tin c tiv e b eh a v io r, d isc u sse s th e p h y sio lo g ic a l mechanisms concerned w ith th e p ro c ess he la b e ls as S tim u lu s f i l t e r i n g . 1 1 "S tim ulus f i l t e r i n g " can ta k e p la c e i n th re e fu n c tio n a l l o c i , ac co rd in g to M arler3 1* The f i l t e r i n g mechanism may be lo c a te d p e r ip h e r a lly , i n th e re c e p to rs (an a lte r n a tiv e alread y r e je c te d above)* 2* The meohanism may be i n a system made up of the r e c e p to rs , a f f e r e n t pathways and the c e n tr a l nervous system as th ey fu n c tio n in normal percep tio n * 128 3* The f i l t e r i n g mechanism may be a s p e c ia l one, a c tiv e to s tim u la tio n by complex s tim u li and "o p e ra tin g only i n c e r ta in c o n te x ts and c o r responding w ith th e o rig in a l L orenzian concept of a 'r e le a s e mechanism*" (M arler, 1961, p . 160). W hile i t i s im possible to c le a r ly decide between th e second and th ird a lte r n a tiv e s provided by M arler, i t i s tem pting to a sc rib e the change brought about through h a b itu a tio n to a change i n th e in n a te r e le a s in g mechanism (IRM) on the grounds th a t th e behaviors in q u e stio n w ith t h i s sp e c ie s have h i s t o r i c a l l y and ty p ic a lly been con sid e re d examples of " re le a s e d b ehaviors" (T in b erg en , 19f>l) • However, th e re i s not any c le a r b a s is fo r r e j e c t in g Marler* s second a lte r n a tiv e In so fa r as we appear to be a long way from u n d ersta n d in g how the c e n tr a l nervous system behaves and fu n c tio n s in normal p ercep tio n # ^ The D is ln h ib itio n H ypothesis The b a sic prem ise o f th e d is ln h lb ltlo n h y p o th e sis, th a t d riv e system s have m u tu ally in h ib itin g r e la tio n s h ip s 2 A lthough now fiv e y e a rs o ld , d e ta ile d review s of v is u a l mechanisms as th ey were understood a t th e tim e of p u b lish in g may be found i n The Handbook of P hysiology, S e c tio n 1 , Volume I , in p ap ers by fe a rtle y , W y , C&*anit and L iv in g sto n . to each o th e r and t h a t changes in s tr e n g th o f one o e n te r w i l l be r e f le c te d by th e appearance o f b eh av io r (FAPs) a p p ro p ria te to th e dom inant c e n te r , i s supported by th e fin d in g s o f t h i s s tu d y . D uring th e sex u al s ta g e p re s e n ta t io n of a m ale stim u lu s alm ost always r e s u l t s i n a dominance o f the a g g re ssio n c e n te r and th is i s r e f le c te d in behavior by a g g re ssiv e re sp o n se s d ire c te d a t th e i n tru d in g m a le . I t was assumed th a t h a b itu a tio n o f a g g re s s iv e re sp o n se s would r e f l e c t a d lm u n ltio n of th e s tr e n g th of the a g g re ssio n by in h ib itin g th e ag g re ssio n c e n te r . T his i s co n sid ered t o be a p ro c e ss analogous to “in t e r n a l in h ib itio n ” in th e F av lo v ian sense* I n t h i s m anner, th e n , th e r e l a t i v e dominance and su b se rv ie n c e of th e r e c ip r o c a l ly i n h i b iti n g d riv e -c e n te rs i s a lte r e d to allow th e se x u a l o e n te r to g a in ascendency even i n th e fa c e o f a stim u lu s , such as a c o n s p e c ific m ale, th a t norm ally e l i c i t s ag g res sion* I n t h i s stu d y th e t e r r i t o r i a l male s tic k le b a c k s th a t were h a b itu a te d to a r e a l male began r a th e r q u ic k ly to respond to th e r e a l male stim u lu s w ith a se x u a l FAP (th e z ig -z a g ) fo llo w in g the most marked d e c lin e in ag g ressio n * F u rth erm o re, t h i s e f f e c t was not stim u lu s s p e c if ic , b u t r a th e r th e in f e r r e d ascendance o f the sex c e n te r was shown by in c re a s e d sex u a l r e s p o n s iv ity toward a fem ale o v er the co u rse o f th e experim ent* 130 I t i s notew orthy th a t t h i s same e f f e c t was n o t found w ith f i s h t h a t were h a b itu a te d to th e model of a male stick leb ack * Here h a b itu a tio n was c le a r ly shown in term s of a decrem ent over days b u t i n only two cases did th e re s id e n t male respond to th e model w ith sex u al be havior* This may w e ll have been because th e model was so weak in i t s a b i l i t y to e l i c i t ag g ressio n t h a t the response re d u cin g e f f e c t o f h a b itu a tio n on th e ag g ressio n c e n te r was in s u f f ic ie n t to depress th e ag g ressio n c e n te r to a low enough le v e l th a t the sex u al d riv e could g a in a s cendance* The in fe re n c e being made by th is d is c u ss io n i s t h a t th e amount o f in h ib itio n r e s u ltin g from h a b itu a tio n to a stim u lu s i s d ir e c tly p ro p o rtio n a l to i t s i n i t i a l a b i l i t y to e l i c i t th e re sp o n se, and a lso th e degree to which th a t resp o n se has been h ab itu ated * T his co n jec tu re p r e d ic ts th a t the g re a te r th e a b i l i t y of a stim u lu s to e l i c i t a g g re ssio n , th e g re a te r w ill be the in h ib itio n o f th e ag g ressio n c e n te r when the a g g r e s s io n - e lic itin g stim u lu s has been h a b itu a te d * I t i s n o t c le a r w hether a response must be h a b itu a te d through c o n sta n t r e p e t it iv e e l i c i t a t i o n o r w hether sim ply r e p e titiv e p re s e n ta tio n of th e stim u lu s w ith o r w ith o u t th e response i s n ecessary or s u f f ic ie n t to b rin g about th e in h ib itio n o f th e d riv e - c e n te r* The answer to t h i s q u e stio n aw aits experim ents 131 on " la te n t h a b itu a tio n ," which would be th e o r e tic a lly and m eth o d o lo g ically analogous to " l a te n t le a rn in g " stu d ie s* R eturning fo r a moment to th e q u e stio n of th e fu n c tio n a l lo cu s of th e e f f e c t of h a b itu a tio n d iscu ssed in th e p rev io u s s e c tio n , we may re fo rm u la te th e q u e stio n i n term s of th e "c e n te r" th e o ry in th e d ls ih h lb itio n h ypothesis* The c e n te r i s a c tiv a te d by two so u rc e s, I n te r n a l and e x te r n a l, each w ith two p o s s ib le p ro c e sse s, e x c ita tio n and in h ib itio n * In s o fa r as th e stim u lu s does n o t change i n t h i s ex perim ent, th e e f f e c t cannot be due to e it h e r an e x c ita to ry or in h ib ito r y p ro cess stemming from any e x te rn a l source* T his leav e s th e locus i n t e r n a l , and th e p ro c e ss, r e s u ltin g i n decreased re s p o n a iv ity r a th e r th an in c re a se d re s p o n a iv ity , in h ib itio n . The E x ten sio n of th e D is in h ib itio n H ypothesis The e x te n sio n of th e d is ln h lb ltlo n h y p o th esis developed i n C hapter I I I p re d ic te d th e occurrence o f displacem ent b eh av io rs as th e r e s u l t of low ering th e le v e l of a g g ressio n r e la tiv e to th e le v e l o f sex in the s itu a tio n where th e male s tic k le b a c k was responding to s tim u la tio n from a male stim u lu s In h is te r r ito r y * This low ering o f a g g re ssio n r e la tiv e to sex was to r e s u l t , a t some tim e in th e course of d ec reasin g th e le v e l of a g g re ssio n , i n a c o n f lic t betw een sex and a g g re ssio n . This c o n f lic t would be m an ifest th e o r e tic a lly by a r e d u ctio n i n th e combined in h ib itio n e x e rte d by b o th the sox and ag g ressio n c e n te rs on th e c e n te rs subsuming n e s t-o o n s tru e tio n and p a re n ta l b e h a v io r. In behavior th is c o n f lic t and subsequent d is ln h lb ltlo n would be r e vealed by the emergence o f n e s t-c o n s tru o tio n beh av io r and p a re n ta l b e h a v io r. As h a b itu a tio n proceeded, th e sexual c e n te r would became dominant as th e s tre n g th of ag g ressio n was f u r th e r depressed and sex would th e n i n h i b i t both n e s t-c o n s tru c tio n and p a re n ta l c e n te rs and the b eh av io rs a p p ro p ria te to th e se c e n te rs would d isap pear* Of te n f i s h (two i n group M M , a l l e ig h t i n group RM ) th a t showed a peak of a g g ressio n and a subsequent peak o f s e x u a lity ( t h i s in c lu d e s one f i s h whose peek of ag g ressio n was masked by what appeared to be i n i t i a l f e a r o f the s tim u lu s ), only s ix showed any displacem ent a c t i v i t i e s . Of th ese s ix , two c le a r ly supported th e h y p o th e sis, two gave q u e stio n a b le support and two gave n eg a tiv e ev id en c e. I t cannot be sa id t h a t th e e x te n sio n of th e d is ln h lb ltlo n h y p o th e sis proposed i n t h i s d is s e r ta tio n wa3 supported but th e r e s u l t s do suggest th a t Improvements i n th e ex p erim en tal method m ight be ap p lied so t h a t th e o p p o rtu n ity fo r a c le a r su p p o rt o r r e f u ta tio n 133 of th e model would he p o ssib le * The m ost im p o rta n t change I n method would be to extend th e o b se rv a tio n s of the m ale stic k le b a c k * a b e h a v io r over a p erio d encom passing more th a n ju s t ag g res s io n p e rio d s d u rin g th e sex u al sta g e to p e rio d s o f c o u rt sh ip and p e rio d s d u rin g which th e r e i s n e ith e r m ale nor fem ale s tim u li p resen t* In th e p re se n t s tu d y , th e p e rio d s of c o n f lic t and displacem ent may have o ccu rred a f t e r th e stim u lu s (m ale) was removed and the r e s id e n t male was no lo n g er b ein g observed* T his assumes t h a t the c o n f lic t could occur w ith o u t th e a g g r e s s io n - e lic itin g stim u lu s b ein g p re se n t* Shis prem ise does n o t seem un re a so n a b le i n t h a t the n europhysio lo g ic a l changes th a t m ight account f o r th e i n h i b iti o n o f th e d riv e c e n te r may be o f a p e rs e v e ra tiv e n a tu re t h a t co n tin u es a f te r th e s tim u la tio n experience* This p o s s i b i l i t y seems l e s s r e mote i f h a b itu a tio n Is viewed as a le a rn in g o r le a r n in g - lik e phenomena. The evidence seems to fa v o r p o s t-e x p e ri e n t i a l p e rs e v e ra tio n and f ix a tio n o f th e meumonlc tra c e in a wide range o f In stru m e n ta l s i t u a t io n s (G lickm an, I 96I)* R eg ard less of what the mechanism m ight b e , modi f ic a ti o n s of th e method used in th e p re s e n t study should fo llo w th e lin e s of in c re a s in g th e d u ra tio n of o b serv a tio n so as to d e te c t any emergence o f d isp lacem en t th a t m ight occur d u rin g c o u rts h ip and th e se x u a l phase i n gen eral as w e ll as d uring p erio d s o f s tim u la tio n from another m ale. I t must be tak en in to account t h a t the p erio d of c o n f lic t migjit be a f l e e tin g in t e r v a l , and th a t th e r a te of change in th e r e la ti v e s tre n g th s o f sex and ag g ressio n due to in h ib itio n v ia h a b itu a tio n of the ag g re ssiv e d riv e would lik e ly vary m arkedly betw een in d iv id u a l fis h * C H A PO !E R V II S U M M A R Y H a b itu a tio n of re p e a te d ly e l i c i t e d resp o n ses i s a phenomenon of g e n e ra lity found w ith many sp e c ie s and w ith a wide range of stim u li# For th e most p a r t , however, the resp o n ses s tu d ie d have b een sim ple re fle x e s # Only in a few s tu d ie s (m ost n o tab le of which were perform ed on b ird s ) were complex, m ulti-com ponent resp o n ses h ab itu ated # The p re s e n t study attem pted to h a b itu a te ag g ressiv e r e sponses of th e t e r r i t o r i a l male th re e -s p in e d stic k le b a c k # The stim u lu s used f o r one group of e ig h t f i s h was a liv e male stio k le b a o k confined in a c le a r p l a s t i c tube# For th e second group of e ig h t f i s h , th e stim u lu s was a crude wooden model of a s tic k le b a c k moved in a c ir c u la r p ath in th e t e r r i t o r i a l m alef s tank# The t e r r i t o r i a l m ales were p re se n te d w ith the a g g r e s s io n - e lic itin g stim u lu s fo r 15 m inutes each day fo r te n days# A th ir d group was also m aintained i n a manner id e n tic a l to the o th er two groups except th ey were given no a g g re ssio n - e l i c i t i n g s tim u la tio n f o r th e f i r s t te n days o f th e experim ent# I n a d d itio n to a g g r e s s io n - e lic itin g s tim u li, a l l f i s h were p re se n te d w ith a g ra v id fem ale i n a p la s tic 135 136 tu b e f o r f iv e m inutes each day th ro u g h o u t th e experim ent i n o rd e r to m easure th e s tre n g th o f th e sex d riv e* Both th e group p re se n te d each day w ith th e r e a l male and th e group p re se n te d w ith th e model male showed a decrem ent In a g g re ssio n over days* B oth stim u li* through d a ily r e p e t i t i o n r e s u lte d In h a b itu a tio n o f ag g re s s iv e resp o n ses* A r e c e n t th e o ry advanced by s e v e ra l e th o lo g is ts a t th e U n iv e rsity o f L eiden p r e d ic ts s p e c if ic changes i n b e h a v io r as th e r e s u l t o f i n h ib it io n of th e M d riv e c e n te r” su b serv in g v a rio u s b e h a v io ra l c a te g o rie s such as sex* a g g re ssio n and p a re n ta l* In s o fa r as sex and a g g re ssio n are th e dom inant b e h a v io rs d u rin g th e phase o f th e r e p ro d u c tiv e c y c le In w hich th e f i s h I n t h i s stu d y were m a in ta in e d , t h i s th e o ry assumes t h a t th e dom inant “ d riv e c e n te rs ” a re sex and aggression* w ith a g g re ssio n b e in g dominant when the e x te r n a l stim u lu s i s a c o n s p e c ific male in tru d in g in to th e t e r r i t o r y o f th e r e s id e n t maldo The b alan ce or dominance r e la tio n s h ip s betw een th e s e two d riv e c e n te rs m ight be u p s e t i f some means were used to I n h ib it one or the oth er* I n t h i s re s e a rc h th e p ro c e ss u n d e rly in g h a b itu a tio n was co n sid ered to be a source of i n t e r n a l i n h ib it io n and was th u s a means o f i n h ib it i n g th e d riv e c e n te r f o r agg ressio n * The e th o lo g ic a l th e o ry th e n p r e d ic ts th a t se x u a l b eh a v io r should emerge as a g g re ssiv e behav io r d e c re a se s in th e p ro c ess of h a b itu a tio n * This p r e d ic tio n was su p p o rted i n two ways by th e group h a b itu a te d to th e r e a l male stim ulus* F i r s t , as a g g re ssio n waned, se x u a l re sp o n se s were observed to th e r e a l male* Second, th e frequency o f sex u al re sp o n se s to th e g ra v id fem ale in c re a se d over th e d u ra tio n of th e ex p erim en t. No such in c re a s e i n se x u a l resp o n se to th e g rav id fem ale was seen w ith th e group of f i s h g iv en no a g g r e s s io n - e lic itin g stim u la tio n * The group t h a t h a b itu a te d to th e model m ale d id n o t show any emergence o f se x u a l b eh a v io r tow ard th e model as a group nor did th ey in c re a s e t h e i r frequency of se x u a l re sp o n se s to th e g ra v id fem ale* The model e l i c i t e d much le s s a g g re ssio n i n i t i a l l y th a n did th e r e a l male* As a r e s u l t (th e o r e t i c a l l y ) th e amount of i n h ib it io n e x e rte d upon th e a g g re ssio n o e n te r through h a b itu a tio n of th e resp o n se to th e model must not have been s u f f i c ie n t to u p s e t or change th e b alan c e of th e d riv e c e n te rs* The p re s e n t w r ite r sought to extend and m odify th e th e o ry p u t f o r t h by th e e th o lo g is ts to p r e d ic t th e o ccu rren ce of displacem ent b eh a v io r i n th e p a r ti c u l a r s i t u a t io n used in t h i s study* T his th e o r e tic a l e x te n sio n was n o t e i t h e r supported nor r e fu te d by th e r e s u lts * S uggestions f o r an improved method of t e s t i n g t h i s ex te n s io n o f th e th eo ry were made on th e b a s is o f th e r e s u l t s of th e p re se n t study* LIST OP REFERENCES LIST OF REFERENCES B aennlnger, R. Waning o f a g g re ssiv e m o tiv a tio n In B e tta s p le n d ln s . Pay chon, S o l. , 1966, ij., 2jj.l-2i(.2. B a rtle y , 3* H. C e n tra l mechanisms of v is io n . I n J , F ie ld (Ed*) Handbook o f p h y sio lo g y ; S e c tio n 1 , N europhyaiology . V o l. i . W ashington, fr.fl.g American p h y s io lo g ic a l S o c ie ty , 19519. P p. 713- 7^0* B artoshuk, A. K. Human n e o n a ta l c a rd ia c a c c e le r a tio n to sound: h a b itu a tio n and d is h a b itu a tio n . Per c e p . Mot. S k i l l s , 1962, l £ , 1, l£ -2 7 . B o y co tt, B* B. L earning I n octopus v u lg a ris and o th e r cephalopoda. P u b l. S ta z . Z o o l. N a p o li., 19!?b, 25, 67-93. ----- B rid g e r, W, H. Sensory h a b itu a tio n and d is c rim in a tio n i n th e human n e o n a te . Amer. J . P s y c h la t., 1961, 117» 991-996. ------------------ ---------- B runt on, T. L . On th e n a tu re o f in h ib itio n and th e a c tio n of drugs upon i t . 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H lnde, R* A* F a c to rs governing th e changes I n s tre n g th o f a p a r t i a l l y In b o rn re sp o n se , as shown by th e mobbing b eh av io r o f th e C haffin ch (F r i n g l l l a c o e le b s ) • I* The n a tu re of th e re sp o n se , and an ex am in atio n of I t s course# Proc# ro y . Soc# B#, 1954-a, 306-331. » F a c to rs governing the changes i n th e s tre n g th of a p a r t i a l l y Inb o rn re sp o n se , as shown by th e mobbing b eh a v io r o f th e C haffinch (F r i n g l l l a c o e le b s ). II* The waning of th e re sp o n se . P ro c. roy* Soo* B. , 1 9 5 ^ * 331-358. • U n ita ry d r iv e s. B rit* j . anlm* B ehav., 1959, T, 130- ll|.l. ~ _________. F a c to rs governing th e changes In th e s tre n g th o f a p a r t i a l l y in b o rn re sp o n se , as shown by th e mobbing b eh a v io r o f th e C haffinch (F r i n g l ll a c o e le b s ) . I I I . 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Oxford, 1951 • • D erived a c t i v i t i e s ; t h e i r c a u s a tio n , b io lo g ic a l s ig n ific a n c e , o r ig in and em ancipation d u rin g e v o lu tio n . Q u art. Rev. B io l. , 1952, 27, 1-32* T inbergen, N ., and l e r s e l , J . A. A. van . D isplacem ent re a c tio n s i n the th re e -sp in e d s tic k le b a c k . Be- h a v io u r, 19^ 7 * 1* 56- 63. T r i p l e t t , N. The e d u c a b ility o f th e p e rc h . Am. J . P sychol. . 1901, 12, 35^-360. Z e ig le r, H. P . D isplacem ent a c tiv ity and m o tiv a tio n a l th e o ry ; a case study in th e h is to r y of e th o lo g y . P sy ch o l. B u ll. , I 96I 4 ., 61, 5 , 362-376. APPENDIX A APPENDIX A SELECTIVE REVIEW OP THE HABITUATION LITERATURE The conceptual-m ethodological shortcom ings of many of th e s tu d ie s of h a b itu a tio n I n th e l i t e r a t u r e su g g est r e s t r i c t i o n of a p h y lo g en etic review o f response decrem ent s tu d ie s to th o se which are more rig o ro u s , w ith prim ary emphasis on in v e s tig a tio n s w ith v e r te b ra te s and h ig h e r I n v e r te b r a te s , more s p e c if io a lly In v e s tig a tio n s s tr e s s in g h a b itu a tio n of more com plicated b eh av io rs as opposed to sim ple r e f le x h a b itu a tio n * F or more com plete, le s s s e le c tiv e rev iew , th e re a d e r i s r e f e r r e d to H a rris (19^3) 9 Thorpe ( 1963)9 and Thompson and Spencer (19 66) *^ In v e rte b ra te s Jennings (1902) stim u la te d S te n to rs and V o r tic e lla w ith a f in e h a ir o r g la s s ro d and was ab le to r e l i a b l y e l i c i t a c o n tra c tio n * Repeated s tim u la tio n s a t a r a t e of one p er second of t h i s s o r t r e s u lte d i n a decrem ent i n th e number o f c o n tra c tio n s e l i c i t e d by th e stim u lu s w ith more and more s tim u la tio n s neo essary b efo re c o n tra c tio n would occur* F u rth e r, he stim u la te d th e organism s w ith „ ^ C ita tio n s i n th e Appendix may be found i n L is t o f R efe ren ce s, page llj.0* 347 a weak w ater c u rre n t* This c o n tra c tio n resp o n se also waned, and more r a p id ly th an to t a o t i l e stim u la tio n * To r u l e out th e p o s s i b i li ty o f fa tig u e as re sp o n sib le fo r th e resp o n se d e f ic it he ad m in istered a NaOl s o lu tio n to th e organism a g re a t number o f tim es and n ever recorded any response d e f ic it d e sp ite the f a c t t h a t th e response to s a l t w ater i s th e same as to t a o t i l e and w ate r c u rre n t stim u latio n * More p re c is e stim u lu s c o n tro l was used w ith th e same organism by D anlsch (1921) i n th a t he used w eights dropped on th e substratum o f th e c o n ta in e r as th e stim ulus* He found a p o s itiv e c o r r e la tio n betw een the amount o f energy used to stim u la te th e organism and the number o f r e p e titio n s n ec essary b e fo re c e s s a tio n of re * sponse* Thus i t appears th a t low I n te n s ity s tim u li h a b itu a te more ra p id ly th an h ig h e r in te n s ity ones* Jennings (190f>) s tu d ie d the e f f e c ts o f drops of w ater on th e su rfa c e of a c o n ta in e r h o ld in g a se a anemone (A lp ta a la a n n u la te )* To th e f i r s t d ro p , th e anemone con* tr a c te d , and sometimes to th e second; to th e th i r d and f o u r th , i t did not* This v a rie d as a fu n c tio n o f th e I n te r v a l betw een d ro p s, becoming a le s s r e l i a b l e phenome non as the i n t e r t r i a l I n te r v a l was extended from one m inute to In te r v a ls o f lo n g er th an f iv e m inutes* P leron (1 9 0 8 ), u sin g m echanical s tim u la tio n , found s im ila r r e s u lts i n o th e r sea anemones (A c tin ia equina and A c tin ia r u b r a ) « The most co n v in cin g evidence f o r h a b itu a tio n i n th e se a anemone comes from th e work o f F lu e re and W alton (1907)* They fe d sm all p ie c e s o f f i l t e r p ap er to se a anemones each day, p la c in g i t on th e same te n ta c le s * W ith in a few d ay s, th e te n ta c le s would no lo n g e r a c c e p t th e f i l t e r paper* T his e f f e c t la s te d s e v e ra l days to a week* Hie tem poral p e r s is te n c e of th e e f f e c t would seem to r u l e out th e e f f e c ts o f fa tig u e * S* J* Holmes (1 9 1 2 ), i n c o n firm a tio n of some e a r l i e r r e s u l t s , was a b le to dem onstrate t h a t the s e a -u rc h in (A rbaola p u n c tu la ta ) would q u ic k ly sto p re sp o n d in g to s u n l lg r t w ith th e u s u a l n e g a tiv e p h o to ta x ic resp o n se w ith re p e a te d p re s e n ta tio n s * That sim ple sen so ry a d a p ta tio n i s n o t s u f f i c ie n t to ex p la in the resp o n se decrem ent i s shown by th e f a c t th a t m e c h a n ic a l-ta c tile s tim u la tio n would r e s to r e th e response* S tu d ie s of resp o n se decrem ent to re p e a te d stim u la tio n i n flatw orm s (p la n a r!a n s ) and a n n e lid s a re review ed by Jacobson (1962)* Worthy o f p a r ti c u l a r m ention i s th e stu d y of th e m arine worm, N ereis p e la g ic a * by C lark (I9 6 0 )* He h a b itu a te d a v a r ie ty of s tim u li: m ech an ical sh o ck s, shadows, l i g h t o f f - s e t and l i g h t o n -set* In g e n e ra l he found t h a t a l l th e se s tim u li h a b itu a te d and t h a t th ey showed a r e l a t i o n betw een I n t e r t r i a l i n t e r v a l and p e r s is te n c e of resp o n se decrem ent* In th e phylum M olluscs we have s e v e r a l c la s s ic l£ o p ap ers th a t are fre q u e n tly quoted* In th e c la s s G astropoda, P ie r on (1913) perform ed an experim ent stu d y in g th e i n t e r stim u lu s in te r v a l as i t a f f e c ts th e response to shading from a b r ig h t l i g h t . He gave each anim al 1 * > shadings la s tin g 1 second. He v a rie d i n t e r t r i a l i n te r v a l between groups between 3 and 120 seco n d s. He found th a t 60 seconds between r e p e titio n s was o p tim al f o r h a b itu a t i o n . T his com parative spacing f o r maximal e ffic ie n c y i n h a b itu a tio n i s an Im portant change from lower sp ec ie s ( e . g . , C lark , 19&0, who found b e s t h a b itu a tio n a t sh o rt In te rs tim u lu s in te r v a ls ) and an emergence o f a p rin c ip le fin d in g in s e v e ra l s tu d ie s o f h ig h e r form s. Another im p o rtan t fin d in g o f p le r o n 's i s th e long term e f f e c ts of th e h a b itu a tio n . He found th a t in te rp o la te d r e s t p e rio d s as long as an hour (h is lo n g e st p erio d ) did not r e in s t a t e re sp o n siv e n e ss. In the land s n a i l , Humphrey ( 1933) found th a t je rk in g o f a board upon which th e s n a il was p laced r e s u lte d i n a w ithdraw al response o f th e an ten n ae. Je rk in g th e board every 2 seconds f o r up to 60 t r i a l s r e s u lte d i n c e s s a tio n of re sp o n se . A fte r a 30 second r e s t p e rio d , th e resp o n se was r e in s t a t e d , b u t few er t r i a l s were needed to b rin g about response ce ssa tio n w ith each s e r ie s o f s tim u la tio n s . The cephalopoda, re p re s e n tin g , w ith t h e i r more advanced sensory equipm ent, th e h ig h e s t c la s s of M olluscs, wore f i r s t in v e s tig a te d i n re fe re n c e to resp o n se d ecre ment by B uytendljk (1933) w ith Octopus v u lg a r is * B uytendljk used a w hite card a tta c h e d to a s t i c k to e l i c i t a w ithdraw al response* F if te e n p re s e n ta tio n s were s u f f i c ie n t to b rin g about th e r e v e r s a l of th e response to one o f " s lig h t in c lin a tio n of th e head towards th e o b je c t* " A fte r m inutes r e s t th e response "spon ta n e o u sly re c o v e rs ," b u t o nly 6 p re s e n ta tio n s a re s u f f i c ie n t to b rin g about h a b itu a tio n * A th ir d $ m inute r e s t p erio d allow s th e resp o n se to re c o v e r b u t only one more p re s e n ta tio n i s n ec essary to b rin g about response c e ssa tio n * A fte r a r e s t p e rio d of one hour i t took. 11 p re se n ta tio n s to no lo n g er evoke response* T his same p a t te r n of h a b itu a tio n , r e s t , reco v ery (b u t few er e l i c i t a tio n s needed fo r re h a b itu a tio n ) i s re p o rte d by G oldsm ith (1927* c ite d by B o y co tt, 195^) f o r th e a tta o k resp o n se o f th e octopus to w ater a g ita tio n * B oycott (19*34-) r e views th ese s tu d ie s and adds confirm ing evidence o f h is own* A review of some o f th e work w ith arth ro p o d s culm in ates th is sh o rt review of work on response d ecre ment w ith in v e rte b ra te s * W ith th e pupae of T enebrlo m o llto r, H o llis (19&3) found th a t re p e a te d e l e c t r i c a l s tim u la tio n a t very low amperage would r e s u l t i n h a b itu a tio n of a c u rlin g resp o n se in 12 to lj?2 stim u la tio n s* ia>2 D is h a b ltu a tlo n w ith t a c t i l e s tim u la tio n seems t o r u le o u t fa tig u e as an e x p la n a tio n o f th e resp o n se decrem ent* D ePorte (1916) s tu d ie d th e waning o f th e d e a th * fe lg n ln g resp o n se i n a w eev il ( Tyohius p l c l r o s t r l a ) Induced by f lip p in g th e organism over on i t s back* He found t h a t . In le s s th an ij.0 e l i c i t a t i o n s , th e resp o n se would no lo n g er occur* The u se o f a new stim u lu s would r e i n s t a t e th e resp o n se ( d is h a b ltu a tlo n ) , b u t few er e l i c i t a t i o n s were needed f o r re h a b itu a tio n * F u r th e r , he found th a t th e decrem ent i n re sp o n siv e n e ss rem ained a f te r 2 hours r e s t* Working w ith s p id e r s , Peckham and Peokham (1887) found t h a t th e re sp o n se to a tu n in g fo rk s tim u la tio n o f th e web was to drop from th e web* They found t h a t w ith re p e a te d e l i c i t a t i o n s t h i s resp o n se would g ra d u a lly d isap p ear* L a te r , more c a r e f u l and q u a n tita tiv e work w ith s p id e r s , w ith v ib r a tio n s o f a more m oderate in * t e n s i t y , has y ie ld e d an im p o rta n t fin d in g * The work o f S zlep ( I 96I 1) w ith Araneus and U loborus c o rro b o ra te s e a r l i e r fin d in g s by D ress (195>2) and F re o h t ( 1958) as to th e e f f e c t of re p e a te d v ib ra to ry and p rey s tim u la tio n s of th e web* With r e p e t i t i o n th e approaoh (p rey c a tc h in g ) resp o n se g ra d u a lly drops o u t, b u t n o t i n a l l or none fa sh io n * The l a s t lin k s o f th e re sp o n s e , th e pouncing tow ards the p re y , d isa p p e a r f i r s t , fo llo w ed by th e locom otor resp o n ses tow ards the p re y and f i n a l l y , oven th e o r ie n ta tio n d isappears* In S a lo p 's stu d y (1961).) th e s tim u li t o be h a b itu a te d were m echanical v ib ra tio n s of th e web r a th e r th a n re sp o n se s to a r e a l prey* The same type of r e s u l t s p re se n ted them selves* The response decrem ent B tarted w ith a slow ing down o f the resp o n se follow ed by appearance of incom plete re sp o n se s, th e n a com plete waning* The im p o rtan t p o in t f o r th is p ro p o sal in th e re se a rc h on s p id e rs i s th e d i f f e r e n t ia l decrem ent of d if f e r e n t components of the response due to re p e a te d e l i c i t a t i o n by a stim u lu s r a th e r than an a l l or none e f f e c t o r sim ple m agnitude an d /o r la te n c y change as i s ty p ic a lly re p o rte d in o th e r in v e rte b r a te h a b itu a tio n stu d ie s* From th e s tu d ie s w ith in v e rte b r a te s we may in d ic a te s e v e ra l im p o rtan t c h a r a c te r is tic s of h a b itu a tio n emerging; 1* At the lo w est l e v e ls , S te n to r and V o r tio e lla , h a b itu a tio n i s obtain ed th a t can be c le a r ly d if f e r e n tia te d from fa tig u e * F u rth e r, a t the same p h y le tlc le v e l, a c le a r s u b s ta n tia tio n of th e g e n e ra l p rin c ip le th a t low in te n s ity s tim u li h a b itu a te more ra p id ly than h ig h e r i n te n s ity ones i s found* 2* W ith the se a anemone, tem poral p e rs is te n c e of lo n g er than m inutes o r hours was found* 1 5 1 + 3* I n M ollusc a i t was dem onstrated th a t d i s t r i b u ted t r i a l s were s u p e rio r to massed t r i a l s i n producing response decrement* When t h i s v a ria b le was in v e s tig a te d a t low er phyla t i c le v e ls the re v e rse was found* i|.* In two s tu d ie s w ith arthropods* th e re i s c le a r evidence f o r d is h a b itu a tio n from new e x te rn a l stim u la tio n * 5* In work w ith spid ers* i t was found th a t h a b itu a tio n of more com plicated response p a tte rn s proceeded n o t in an a l l or none fashion* b u t w ith d if f e r e n t components drop p in g out sooner th a n o th ers* V e rte b ra te s Thorpe (1963) s t a t e s th a t " th e re has as y e t been no c r i t i c a l work on h a b itu a tio n i n fis h e s * • • *" He c it e s se v e ra l re fe re n c e s o f in fo rm al o b serv a tio n s such as tam ing of f i s h i n aquaria* b u t re p o rts one stu d y under h is se c tio n heading of T ria l-a n d -e rr o r L earning th a t i s d ir e c tly re le v a n t to th e experim ent u n d ertak en i n th is d is s e rta tio n * and i s c e r ta in ly evidence f o r h a b itu a tio n of ag g ressiv e behavior* T r i p l e t t (1901) found th a t p erch would cease " a tta c k in g " minnows th a t were p ro te c te d behind a g la ss p la te * However* th e p o s s ib i li ty of 155 n e g a tiv e rein fo rcem en t e f f e c ts of c o n fro n tin g th e g la s s b a r r ie r a t g re a te r th a n n e a r zero v e lo c ity may in d ic a te th e re a so n f o r T&orpets in c lu s io n of th i s stu d y i n h is T ria l-a n d -e rro r L earning se c tio n * Baexminger ( 1966) has p u b lish ed a paper p u rp o rtin g to dem onstrate th e waning o f ag g ressiv e m o tiv a tio n i n B eta s p le n d in s * This stu d y was done i n such a manner th a t a b e ta was provided w ith a ta n k a t one end o f which was a model of an o th er male b e ta , and a t th e o th e r end was an o th er ta n k w ith a liv e male b eta* I n the c e n te r was an a re a where th e ex p erim en tal su b je c t could e n te r and not view e i th e r stim ulus* The m easure o f ag g ressiv e m o tiv a tio n was th e amount o f tim e sp en t i n th e c e n te r a re a in a sin g le 16 hour sessio n * The ex p e rim e n ta l f i s h spent more tim e i n the c e n te r compartment o f th e apparatus as tim e went on* This was in te r p r e te d to mean th a t th e f i s h was no lo n g er m otivated to view e i th e r stim u lu s because h is ag g ressiv e m o tiv a tio n had waned* An e q u a lly v a lid and e q u a lly re aso n ab le i n te r p r e ta ti o n was th a t th e ex p erim en tal f i s h was avoiding the s tim u li out of f e a r r a th e r th an o u t of lowered a g g re ssiv e m otiva tio n * T h is study cannot be co n sid ered a c le a r dem onstra tio n of h a b itu a tio n of a g g re ssio n , alth o u g h i t i s sug g e stiv e o f such a p o s s ib ility * R ecen tly a study was u n d ertak en i n th e U n iv e rsity U> 6 „ o f S outhern C a lifo rn ia la b o ra to ry by E« J* Wyers* M * J* H ers and H* V. S . Peeke (unpublished) In w hich t e r r i t o r i a l male s tic k le b a c k s were p re se n te d w ith f iv e m odels* each p re se n te d f o r two m inutes eaoh day over a p e rio d ran g in g from lit to 36 days« The models v a rie d only In amount o f red 00lo r on th e a n te r io r v e n tr a l s u rfa c e * ra n g in g from no re d c o lo r to a model t h a t was e n t ir e l y red* P rev io u s s tu d ie s had shown th e p a tte r n of re d c o lo r to be im p o rtan t as a r e le a s in g stim u lu s f o r ag g ressiv e resp o n d in g i n th e s tic k le b a c k (Tinbergen* 1951)* T his study re v e a le d th a t a decrem ent I n resp o n d in g to a l l models r e s u lte d over days* S e v e ra l v a ria b le s c o n trib u te d to more u n c o n tro lle d v aria n ce th a n was n e c e ssa ry I n suoh a study* b u t th e r e s u l t s do in d ic a te th a t* a t l e a s t to one s p e c if ic shape and s iz e o f model* a g g re ssiv e behavior* as Indexed by th e number o f b ite s a t th e model* does _ wane* F u rth er* o r ie n ta tio n s toward th e models did n o t seem t o wane* F ra n z is k e t (1955* c ite d i n Thorpe* 19&3) found w ith s p in a l fro g s th a t re p e a te d cutaneous s tim u la tio n brought about b o th stim u lu s s p e c if ic and resp o n se s p e c if ic h a b itu a tio n of th e w iping response o f th e hind leg* The evidence on work w ith r e p ti l e s i s alm ost as sc a rc e as th a t w ith amphibia* Humphrey (1933)> u sin g m echanical s tim u la tio n ap p lied to th e s h e l l o f th e 157 t u r t l e , found t h a t ta p s p re se n ted a t two second in te r v a ls took only 7 p re s e n ta tio n s t o b rin g about c e s s a tio n o f th e limb w ithdraw al response* Humphrey c a r r ie d th e work one s te p f u r th e r ; he s tru c k th e s h e ll very hard n ea r th e loous o f th e ta p s and was ab le to dem onstrate d ls h a b itu a - tio n by so doing (l* e « , ta p s o f th e hammer a t th e o r ig in a l lower i n t e n s i ty were now ab le to e l i c i t a resp o n se)* R epeated d is h a b ltu a tlo n f a ile d to r e - e s ta b li s h th e r e sponse* T his may in d ic a te a sub-zero h a b itu a tio n phenomenon w herein th e response to th e low i n t e n s i ty ta p s i s so com pletely h a b itu a te d th a t i t I s no lo n g er p o s s ib le to in d ic a te a resu rg en ce a f te r a d is h a b ltu a tlo n a tira u lu s« She work w ith b ir d s , w hile i n q u a n tity n o t much g re a te r th an w ith r e p t i l e s and am phibians, i s accen tu ated by one p a r ti c u l a r a u th o r1 s work (H inde), to such an ex te n t t h a t th e c o n trib u tio n from t h i s p h y le tlc le v e l dom inates th e work on resp o n se decrement* Because th e work of Hinde w ith th e C haffinch ( 19* > l{ .a , bs 19^ 1) i s c e n tr a l to th e m ethodological con c e p tu a l framework of t h i s re s e a rc h , i t m ight be w e ll to summarize th e experim ents i n some d e ta il* Hinde (19f3f>*) i s concerned w ith th re e prim ary p o in ts * (a ) a d e s c rip tiv e a n a ly s is o f th e mobbing re sp o n se , (b) an a n a ly s is o f some o f th e re le a s in g s tim u li f o r th e mobbing re sp o n se , and (c ) some experim ents d ire c te d a t th e d em onstration of th e in b o rn n a tu re of th e response* The s u b je c ts w ere ch af fin c h e s 5 u s u a lly w ild caught (e x c e p t where n o te d below) and m ain tain ed in screen ed a v ia rie s * The mobbing resp o n se made i n th e fa c e of a p re d a to r (ow l, sn ak e, e tc * ) i s c h a ra c te riz e d by th e ch a ffin ch es* fa c in g th e p re d a to r w ith c r e s t r a is e d , i t s leg s s l i g h t l y fle x e d , th e wings s llg jh tly r a is e d and a b o d ily je rk in g from s id e to sid e accompanied by the c a l l "ch in k chink*" The h ig h e r th e r a t e o f c a ll i n g , th e g r e a te r th e number of n o te s in each phrase* The r a te and freq u en cy of c a ll i n g i s p o s itiv e ly c o r re la te d w ith the s tr e n g th o f the mobbing resp o n se and i t i s used as th e m easure of mobbing* In re g a rd to th e r e le a s in g s tim u li f o r mobbing, H a rtle y , 19*>0 ( c ite d by H inde, 19^l{.a and H inde, 19£L{.b) show d iffe re n c e s between s tu f f e d owls and wooden models p o sse ssin g v a rio u s components o f "ow lness*" C ontour, p a tte r n of c o lo rs and su g g e stio n o f owl and beak a re im p o rtan t v a r ia b le s and d e v ia tio n s from th e se c h a r a c te r is tic s r e s u l t i n lowered re sp o n siv e n e ss to th e stim u li* C le a rly th e s tu f f e d owls and th e o th e r n a tu r a l p re d a to rs such as a s tu ffe d s t o a t and a liv e g ra s s snake were ab le to r e li a b l y e l i c i t th e mobbing response* Hinde* s d a ta from h a n d -ra is e d b ird s gave evidence of th e In n a te n e ss o f th e response* The r e s u l t s were c o n s is te n t w ith an in n a te i n t e r p r e ta ti o n , w ith a m a tu ra tio n a l determ inant o f emergence o f th e response I n a complete form* For th e purpose o f the p re se n t re s e a rc h , th e most im portant fin d in g In t h i s study was th e waning o f the resp o n se over a p erio d o f 30 m inutes of c o n sta n t stim u la tio n w ith e ith e r a s tu ffe d owl o r a model* The course o f th is waning e f f e c t commenced w ith a low le v e l of c a llin g th a t ra p id ly In creased to a peak w ith in about 3 to 6 m inutes* This i n i t i a l In c re a se to a peak le v e l l a c a lle d the "warm-up e f fe c t" (H inde, 195>lt>a, b)* A fte r th e peak i s re a c h e d , th e response slow ly d ecreases u n t i l i t re ach es a low le v e l o r ceases a lto g e th e r* The d isco v ery of t h i s response waning to a p re d a to r stim u lu s le a d s to a study (H inde, 195l|ft) e n tir e ly devoted to th e in v e s tig a tio n of the waning o f t h i s resp o n se and to th e f a c to r s th a t in flu e n c e th i s waning* There a re fo u r Im portant m ajor p o in ts exposed in t h i s stu d y : a* By p re s e n tin g a liv e owl or owl model fo r 30 m inutes In one day, th e n in te r p o la tin g v arious r e s t in te r v a ls b efo re a p re s e n ta tio n o f the same stim u lu s on th e n ex t day, and ta k in g a m easure of th e number o f c a ll s g iv en during th e f i r s t s ix m inutes on b o th occasio n s one may c a lc u la te th e per c e n t d ecrease from day 1 to day 2 In th e r a te of c a llin g * This re p re s e n ts a m easure o f re sp o n se re c o v e ry as a fu n c tio n o f In te r p o la te d r e s t* Hinde found t h a t , up to 30 m in u tes, th e amount of re c o v e ry from th e f i r s t day to th e second was p o s itiv e ly c o rre la te d w ith I n te r p o la te d tim e , re a c h in g an asym ptote o f about $0% re c o v e ry a f t e r a r e s t I n te r v a l o f 30 m inutes* R e st I n te r v a ls lo n g e r th a n 30 m in u tes had no f u r th e r a p p re c ia b le e f f e c t on th e re c o v e ry . T his l a s t f a c t demon s t r a t e s th e e x is te n c e o f a lo n g -term e f f e c t th a t does n o t re c o v e r betw een days* M uscular and sen so ry f a tig u e i s ru le d o u t i n b o th th e s h o rt-te rm and lo n g -term p ro c e ss by o b s e rv a tio n o f th e c h a ffin c h u sin g th e sane m uscles i n o th e r a c t i v i t i e s lim nedlately a f t e r c e s s a tio n o f th e stim u lu s* C h affinch es were p re se n te d w ith a l i v e owl in t h e i r In d iv id u a l a v ia r ie s f o r 20 m in u tes on each of tw elve days* Hinde found t h a t th e average re sp o n se to th e owl decreased from day to day in a n e g a tiv e ly a c c e le ra te d manner* k s in g le p r e s e n ta tio n o f 20 m inutes was n o t as e f f e c tiv e a s d a ily t r i a l s of 3 * K* O I> £ m inutes re p e a te d ov er days i n re d u c in g th e re sp o n se ra te * This i s a m assed v e rsu s d is tr ib u te d 161 e f f e c t d em onstrating th e g re a te r e ffe c tiv e n e s s of spacing of t r i a l s * In com paring th e two waning p ro c e sse s; th e sh o rt-term * resp o n se s p e c if ic decrem ent, and th e long-term * stim u lu s s p e c if ic component (h a b itu a tio n )* th e s h o rt term p ro cess i s c h a ra c te riz e d by r e l a ti v e l y ra p id recovery* n o n -stim u lu s s p e c if ic ity (i*e** th e resp o n se i s depressed f o r a l l s tim u li t h a t e l i c i t th a t response* n o t ju s t to th e stim u lu s t h a t was used to b rin g about the response decrem ent) * and massed p re s e n ta tio n s are more e f f e c tiv e th a n d is trib u te d * In th e case o f long-term * stim u lu s s p e c if ic decrem ent (h a b itu a tio n )* reco v ery i s very slow* response d ecre ment to one stim u lu s i s shared by o th e r s tim u li o nly to th e e x te n t th a t th ey have common c h a r a c te r is tic s (a stim u lu s gene re a liz a tio n -lik e phenomenon)» and spaced p re s e n ta tio n s of th e stim u lu s are more e f f e c tiv e th a n massed* I n a l a t e r study* Hinde (19&1) showed t h a t s e v e ra l c h a r a c te r is tic s of th e resp o n se wane a t d if f e r e n t r a te s * That is* n o t a l l components o f th e complex behavior* mobbing* drop out o r a re reduced in frequency a t th e same ra te * This i s re m in isc e n t of th e same phenomenon found by S zlep in s p id e rs and su g g ests th a t t h i s c h a r a c te r is tic may be c e n tr a l to th e p ro o ess o f h a b itu a tio n o f b eh av io rs th a t are of th e c h o ln e d -re fle x # f lx e d - a c tio n - p a tte r n v a r ie ty as compared w ith th e sim p ler r e f le x iv e h a b ltu a - tio n s t h a t make up th e su b stan ce o f most o f th e p u b lish ed work« Another d em o n stratio n of d i f f e r e n t i a l r a te s o f response component h a b itu a tio n i s found i n an a r t i c l e by Melsack# e t al* (1961) where i t was dem onstrated th a t th e p re s e n ta tio n o f th e hawk-shaped s ilh o u e tte flow n over th e to p of is o la te d cages d u rin g th e f i r s t 20 days of l i f e of young ducks r e s u l t s i n h a b itu a tio n o f th e norm al f e a r r e sponse shown by n aive duck to t h i s fig u re * A r e a n a ly s is of th e o rig in a l p ro to c o ls in th a t experim ent by Melsaok (19^1) re v e a le d a c r i t i c a l phenomenon* I t was n o t th a t th e ducks no lo n g er p aid a tte n tio n to th e hawk shape# but r a th e r th a t th e y co n tin u ed to o r ie n t toward th e stim ulus# w hile th e fe a r resp o n se had been h a b itu a te d # Examples of h a b itu a tio n w ith mammals (in c lu d in g humans) i s more abundant th an a t any o th e r p h y le tlc le v e l b u t i s confined to h a b itu a tio n o f r e f le x type b eh av io rs r a th e r th an complex ch ain s or fix e d p a tte rn s * W ith ra ts # G r i f f i th (1920, c ite d in H a r r is , 19^3) stu d y in g p o s t- r o t a ti o n a l nystagmus# found th a t r o ta tio n a t lt5 rpm p ro duced c e s s a tio n o f p o s t- r o ta tio n a l nystagmus i n between 12 and 19 days* O cular movements were s im ila r ly reduced* 163 U sing r a b b it s , Maxwell, Burke, and Re3to n (1922) found r e s u l t s s im ila r t o G r i f f i th (1920) and f u r th e r , demon s tr a te d t h a t n e ith e r fa tig u e nor sensory a d a p ta tio n were re sp o n sib le f o r th e observed e f f e c ts by producing the same nystagmus resp o n se a t o r ig in a l h ig h le v e ls by a u ra l ir r ig a tio n * P ro sse r and H unter (193 6 ), observing b o th th e le g movement s t a r t l e resp o n se and th e b u rs t d isch arg es from th e gastrocnem ius m uscle t o a u d ito ry c lic k s , found th a t th e s t a r t l e response d isap p ea rs w ith a dim in u tio n o f th e amount o f m otor d isch arg e and th e len g th o f a f t e r - d i s charge* A fte r £ to 30 m inutes r e s t , th e r e f le x was again e l i c i t e d by th e c l i c k stim u lu s, b u t few er p re s e n ta tio n s were n ecessary to re h a b itu a te * I t was also found th a t a pinch on the t a i l (a d is h a b ltu a tlo n stim u lu s) would b rin g back resp o n siv en ess* P ro sse r and H unter extended t h e i r study to se v e ra l o th e r re fle x e s i n th e r a t s th e le g and t a i l je rk response to a tap on th e t a l l , th e same response to a ta p on th e back, th e resp o n se to e l e c tr i c s tim u la tio n on th e t a i l , th e c ro s s -le g r e f le x to e l e c t r i c a l stim u la tio n of th e saphenous nerve* They found th a t resp o n siv en ess could be r e - i n s t i l l e d by d is h a b ltu a tlo n in a l l th e o ases cite d * Lehner ( 19^ 1) s tu d ie d , among o th e r re fle x e s i n o th e r organism s, th e r e s p ir a to r y s t a r t l e response of th e r a t to a u d ito ry buzzer stim u la tio n * she r e s p ir a to r y response dim inished and d isa p p e a re d , b u t was r e in s ta te d by d is - h a b itu a tio n produced (as by P ro sser and H unter, above) ; by a pinch on the t a i l . Subsequent re h a b ltu a tio n s and d is h a b itu a tio n s r e s u lte d i n le s s and le s s re co v ery a f t e r the t a i l pinches and few er and fewer t r i a l s to re h a b itu a te H a rris (19l}-2) used e l e c t r i c a l s tim u la tio n ap p lied te n tim es p e r day ov er a number of days and found a ty p ic a lly n e g a tiv e ly a c c e le ra te d curve which, however, never reached a le v e l o f no response* B ridger ( 1961)» stu d y in g the c a rd ia c and gross movement responses o f human neonates t o to n e s, a stream of a i r , and h o t and co ld w a te r, found d im in u tio n of re * sponse w ith r e p e t it i v e p re se n ta tio n s* B artoshuk ( I 962) p re se n te d to n es and p a tte r n s o f tones to human neonates and measured c a rd ia c a c c e le ra tio n . He found t h a t h a b itu a tio n occurred and could be d ish a b itu a te d * By u sin g a p a t te r n o f tones h a b itu a tio n a g a in appeared, end t h i s a lso could be d ish a b itu a te d * The d is h a b ltu a tlo n stim u lu s in th is case was a r e v e r s a l of th e tone p a tte rn s * This i s good evidence fo r a stim u lu s s p e c if ic it y of th e stim u lu s h ab itu ated * S tu d ie s w ith a d u lt humans, as w ith a ll th e s tu d ie s c ite d h ere w ith mammals, concern sim ple and r e f l e x r e sponses* W ith th e ey eb lin k response to au d ito ry stlm u la - tio n O ld fie ld (1937) found th e c h a r a c te r is tic fin d in g of j e a s ie r h a b itu a tio n v i t h m ild* j u s t su p ra -th re sh o ld s tim u li compared to more in te n s e sounds, although b o th oould be h ab itu ated * F u rth e r, he found t h a t d is h a b ltu a tlo n was more e a s ily dem onstrable w ith h a b itu a te d h ig h in te n s ity stim u li* Using th e g alv an ic s h in response to a u d ito ry s tim u li, Coombs (1938) found t h a t to n es o f a l l I n t e n s i t i e s eould be h a b itu a te d , and th e ty p ic a l n e g a tiv e ly a c c e le r ated curve of response was obtained* Sokolov and p a ra - monova ( 19& 1) dem onstrated h a b itu a tio n of th e o rie n tin g response (d u ra tio n of GSR) to a 1000 cps to n e of 70 db* F u rth e r, th e se au th o rs showed a good g e n e ra lis a tio n curve j I resp o n se to *> 00 and 2000 cps was dim inished due to h a b itu a tio n of th e 1000 ops to n e , and 2£0, I 4 .0OO, and 8000 ops to n es were le s s a ffe c te d by th e p rio r h a b itu a tio n to the 1000 cps tone* S his study was very b ad ly re p o rte d and f u r th e r a n a ly sis o f th e r e s u l t s i s im possibleo Another e n tir e a re a o f human and anim al h a b itu a tio n th a t has j u s t been touched upon in t h i s review and w ill n o t be review ed because i t adds l i t t l e to what h as alread y been dem onstrated i n the s tu d ie s c ite d , I s the a r e a o f human nystagmus* S u ffic e to say th e nystagm us from both a u ra l i r r i g a t i o n and r o ta tio n h a b itu a te s r a th e r re a d ily * From th e s tu d ie s w ith v e r te b ra te s added to th e work c ite d on In v e rte b ra te s i n th i s re v ie w , we may d e sc rib e soma g e n e ra l phenomenon and c h a r a c te r is tic s o f h a b itu a tio n , These co n clu sio n s must be considered most te n ta tiv e and should be tak en only as a p re lim in a ry f i r s t ste p toward in te g r a tin g a v ery d iv e rs e and s c a tte r e d l i t e r a tu re* (For a d isc u ssio n of h a b itu a tio n and neuronal sub s t r a t e s th e re a d e r i s r e f e r r e d to Thompson and Spenser*s (1966) paper devoted e n tir e ly t o th e su b je c t* ) 1* One study w ith humans and one w ith b ird s sub s ta n tia te d th e g e n e ra l p r in c ip le o f low er in te n s ity s tim u li b ein g more e a s ily h a b itu a te d th a n h ig h e r i n te n s ity s tim u li th a t was dem onstrable a t th e low est p h y le tic le v e l stu d ied * 2 . Temporal p e rsiste n c e * e s p e c ia lly w ith b ird s and f i s h u t i l i z i n g more com plicated response p a tte rn s * was dem onstrated over p e rio d s in excess of 2^ hours and p r io r e f f e c ts o f each day compound over days up to th e p o in t of asym ptote of response decrement* 3* Work w ith b ird s showed the s u p e r io r ity of spaces v ersu s massed p ra c tic e th a t was demon s tr a te d w ith h ig h er in v e rte b r a te s b u t not w ith lower (a n n e lid s)* 1 } .* D is h a b ltu a tlo n e f f e c ts were dem onstrated i n two s tu d ie s o f s t a r t l e response h a b itu a tio n w ith 167 r a t s end i n a stu d y w ith t u r t l e s su p p o rtin g and ex ten d in g t h i s fin d in g w ith in v e r te b r a te s . ; 5 > * With b ird s i t was shown t h a t when a complex resp o n se i s s tu d ie d , t h a t h a b itu a tio n proceeds n o t i n an a l l o r none fa sh io n b u t w ith d i f f e r e n t components o f th e o r ig in a l resp o n se drop p in g o u t a t d if f e r e n t r a t e s . This su p p o rts th e fin d in g w ith s p id e rs a lso u t i l i z i n g a more E com plicated response* F urth er* ag ain w ith b ird s i t was found t h a t w ith a stim u lu s th a t o r d in a r ily evokes escape behavior* th a t th e escape response h a b itu a te d b u t th e o r ie n ta tio n component did n o t. Of th e fiv e g en e ral phenomena and c h a r a c te r is tic s of h a b itu a tio n which have been tra c e d through t h i s phy- l e t i c review o f th e lite r a tu r e * th re e are in c o rp o ra te d i n t h i s s tu d y . These a re : 1 . Temporal p e rsiste n c e * which seems th e most c e n tr a l phenomenon c h a r a c te r is tic of h a b itu a ti o n as opposed to o th e r d ecrem ental p ro c ess such as fa tig u e and sen so ry a d a p ta tio n . 2 . S tim ulus I n te n s ity and r a t e of h a b itu a tio n seemed to be a n e a rly u n iv e rs a l fin d in g ( the h ig h er th e in te n s ity th e slow er th e h a b itu a tio n ) b u t had n o t been stu d ie d w ith fix e d a c tio n p a tte r n h a b itu a tio n * D if f e r e n tia l r a te s of h a b itu a tio n o f oomponents o f a b eh av io r c la s s were o f I n t e r e s t because th e s e components may be ordered on a sub je c tiv e b a s is as being more or le s s in te n s e e x p re ssio n s o f a g iv en m otivation* For exam ple, ag g ressiv e responses of h ig h i n t e n s i ty m ight be h a b itu a te d a t a f a s t e r r a t e th a n le s s In te n se ex p ressio n s o f aggression* T h is c h a r a c te r is tic was a lso stu d ie d because i t i s a unique a ttr ib u te of h a b itu a tio n of complex b eh av io r p a tte rn s as opposed to r e f le x h a b itu a tio n * APPENDIX B APH3NDIX B DESCRIPTIONS O P BEHAVIORS RECO RD ED A N D THEIR DEFINING CHARACTERISTICS N e st-C o n a tru c tio n Sand p ic k in g — -Head down, t a i l up p o s tu re , f i s h sucks m a te ria l in to m outh, re tu r n s to h o riz o n ta l and s p i t s o u t m a te ria l* Frequency of 1 f o r e&oh o c c u rre n c e • P ushing— »Head down, t a i l up p o stu re approaching o r a t ta in in g v e r t i c a ll t y * Pushes w ith sn o u t a g a in s t n e s t s e v e r a l tim es i n ra p id succession* Frequency o f 1 f o r each "b o u t* ” (ttB outtt d efined as a com p le tio n o f a sh o rt s e rie s o f p u sh es, te rm in a te d by a w ith d raw al from th e n e st and assum ption of a changed p o s tu re .) B oring—H o riz o n ta l or n e a r h o riz o n ta l p o stu re w ith sn o u t p re sse d i n t o n e st entrance* Frequency of 1 f o r ea ch sn o u t in je c tio n in p ro p e r p o stu re* G luing—F is h assumes h o riz o n ta l p o stu re w ith t a i l e le v a te d a t base and locom otes slow ly over n e s t w ith c o n ta c t on n est* D uring t h i s movement the f i s h s e c r e te s a s tic k y su b stan ce from h is kidney* Frequency of 1 f o r each com plete occurrence* M a te ria l t e s t i n g —R epeatedly p io k s up and s p i t s o u t m a te ria l from bottom , th e su rfa c e o r suspended i n w ater* Frequency of 1 f o r each p ie c e o f m a te ria l te s te d and e i t h e r r e je c te d or used* C r e e p in g th ro u g h --F is h s t a r t s t h i s b ehavior by p la c in g h is snout i n n e s t e n tra n ce (as i n "boring" above) and co n tin u es to push h im self i n to n e s t and, by back and f o r th "squirm ing” movements, p ro p e ls h im se lf through th e n est* This sometimes tak es s e v e ra l seconds* Frequency of 1 f o r each occur rence* A ggression Charge—W ith d o rsa l sp in e s e re c te d , th e f i s h swims r a p id ly d ir e c tly a t the stim ulus* The charge i s i n i t i a t e d a t a d is ta n c e o f a t l e a s t one body length* Each charge co u n ts a frequency o f ! • C ontact w ith stim u lu s o r stim ulus tube i s n o t a c r ite r io n * B ite —F ish c o n ta c ts stim u lu s o r stim u lu s tu b e w ith mouth* Each mouth c o n ta c t w ill count as a f r e quency of 1* T h reat—F is h stands w ith head down, t a l l up p o stu re f r e q u e n tly w ith broad sid e tow ards th e stim ulus* DorsaX f i n s and sp in es erected * Each occurrence cou n ts a frequency o f 1* Z ig -zag —While moving tow ards th e stim u lu s, th e f i s h makes a sidew ays leap follow ed hy a jump d ir e c tly toward the stim ulus* Each com bination o f a s id e ways th en forw ard ra p id movement w il l count as a frequency o f 1 zig-zag* Leading—U sually a f te r a s e r ie s of z ig -z a g s * th e male w ill swim smoothly* w ith o u t h a ltin g to th e n e s t entrance* Each occurrence counts a frequency of 1 * P a re n ta l Panning—The f i s h stan d s a t th e en tra n ce o f th e nest* i t s snout p o in tin g a t th e entrance* The body i s o rie n te d w ith th e n e s t e n tra n ce w ith a v e r ti c a l p lan e 30° from h o riz o n ta l* The t a i l and h in d p e rt o f th e f i s h move back and f o r th as in the normal manner to p ro p e l th e f i s h forw ard* w hile th e p e c to r a l f in s move in a manner norm ally used to p ro p e l th e f i s h backward* The two a n ta g o n is tic movements c a n c e l each o th e r out and th e f i s h r e m ains s ta tio n a ry * b u t d ir e o ts a stream o f w ater forw ard* Bach occurrence o f a bout o f fan n in g counts a frequency o f 1* D epression o f th e pen d e fle c tio n b u tto n f o r th e d u ra tio n of th e fan n in g bout w i ll re c o rd a d u ra tio n measure also* O ther B ehavior O rient a t ! on—Each o r ie n ta tio n of th e f i s h toward th e stim u lu s th a t I s preceded by a d i s t in c t p o s tu ra l change counts as a frequency of 1 o rie n ta tio n *

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Asset Metadata

Creator Peeke, Harman Van Slyck (author)

Core Title Habituation Of Aggression In The Three-Spined Stickleback (Gasterosteus Aculeatus L.)

Degree Doctor of Philosophy

Degree Program Psychology

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag OAI-PMH Harvest,psychology, experimental

Language English

Contributor Digitized by ProQuest (provenance)

Advisor Wyers, Everett J. (committee chair), Savage, Jay Mathers (committee member), Slucki, Henry (committee member)

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c18-113609

Unique identifier UC11359960

Identifier 6700419.pdf (filename),usctheses-c18-113609 (legacy record id)

Legacy Identifier 6700419.pdf

Dmrecord 113609

Document Type Dissertation

Rights Peeke, Harman Van Slyck

Type texts

Source University of Southern California (contributing entity), University of Southern California Dissertations and Theses (collection)

Access Conditions The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the au...

Repository Name University of Southern California Digital Library

Repository Location USC Digital Library, University of Southern California, University Park Campus, Los Angeles, California 90089, USA