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Septal lesions and emotionality in the rat Wexler, Norman 1970

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SEPTAL  LESIONS  AND  IN T H E  EMOTIONALITY  RAT  by  NORMAN B . S c ,  A thesis  submitted in  partial  WEXLER  McGill  University,  to  Faculty of  the  fulfillment Master  in  the  of  of  1967  Graduate  the  degree  Studies, of  Arts  Department of  Psychology  We a c c e p t t h i s required  thesis  as  conforming  to  standard  THE U N I V E R S I T Y  OF  BRITISH  September,  1970  COLUMBIA  the  In  presenting  an  advanced  the I  Library  further  for  his  of  this  thesis  degree shall  agree  scholarly  by  this  at  the University  make  that  it  purposes  written  for  financial  of  September  gain  Columbia  3 D , 197.0  of  Columbia,  British  by  for  shall  that  not  the requirements  reference copying  t h e Head  i s understood  Pgyrhnlngy  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, Canada  of  for extensive  may b e g r a n t e d It  fulfilment  available  permission.  Department  Date  freely  permission  representatives. thesis  in p a r t i a l  of  I agree and this  be a l l o w e d  or  that  study. thesis  o f my D e p a r t m e n t  copying  for  or  publication  without  my  ACKNOWLEDGEMENTS  Thanks to Dr. Rod Wong and Dr. D.J. A l b e r t f o r bearing with t h i s thesis.  John Jamieson rendered s t a t i s t i c a l  advice and the U n i v e r s i t y of B r i t i s h Columbia Health Service k i n d l y provided laboratory facilities.  ii  ABSTRACT  The e f f e c t s of s e p t a l l e s i o n s i n r a t s on the hyperglycaemic response to s t r e s s , adrenal weight and water intake were i n v e s t i g a t e d . times p o s t - o p e r a t i v e l y  Tested four  at approximately weekly i n t e r v a l s , s e p t a l r a t s d i d  not manifest s i g n i f i c a n t l y d i f f e r e n t r e s t i n g blood glucose l e v e l s than c o n t r o l animals nor d i d they demonstrate an abnormal degree of hyperglycaemia f o l l o w i n g periods of unavoidable foot shock. was evident i n s e p t a l subjects compared to c o n t r o l s . was s i g n i f i c a n t l y higher among s e p t a l s .  No adrenal hypertrophy D a i l y water intake  The r e s u l t s concerning blood sugar  l e v e l s and adrenal weights are taken as evidence that s e p t a l rage may not represent true hyperemotionality  since c e r t a i n appropriate  physiological  concomitants are absent. A second experiment i n v e s t i g a t e d r e s i s t a n c e to capture and handling and aspects of open f i e l d behaviour i n s e p t a l and c o n t r o l r a t s f o l l o w i n g i n j e c t i o n s of chlorpromazine hydrochloride  or s a l i n e .  Septalectomized r a t s  r e s i s t e d handling and capture to a greater extent than c o n t r o l s ,  traversed  fewer squares and demonstrated l e s s rears i n an open f i e l d , and had a greater tendency to leave a home cage and enter an open f i e l d .  Chlor-  promazine a f f e c t e d n e i t h e r c o n t r o l nor s e p t a l subjects except to suppress the tendency of s e p t a l r a t s to leave a home cage and enter an open f i e l d . The behaviour of s e p t a l r a t s i s discussed emotionality.  iii  i n terms of apparent  TABLE OF CONTENTS  Acknowledgements  i i  Abstract  —  Introduction  •  ;  •  1  Experiment I  6 Subjects  6  S u r g e r y and H i s t o l o g y —  6  Apparatus  7  Procedure  7  Results  9 15  Experiment I I Introduction  :  15  —  17  Subjects  •— 1 7  S u r g e r y and H i s t o l o g y Apparatus  •  and C o n c l u s i o n s  Bibliography Appendix A  17 19  Results Discussion  i l l  •  32 38  •  4-3  iv  INTRODUCTION  The past f i f t e e n years have seen a large number of studies the e f f e c t s of septal forebrain damage.  concerning  Some of these e f f e c t s , known  c o l l e c t i v e l y as the septal syndrome are; (a) increased water intake (Blass and Hanson, 1970); (b) both p o s i t i v e and negative changes i n general a c t i v i t y l e v e l s depending on the component behaviours Raphaelson, 1966;  measured, (Douglas and  Nielson, Mclver, and Boswell, 1965); (c) f a c i l i t a t i o n of  a preoperatively learned conditioned two-way active avoidance response and of postoperative a c q u i s i t i o n of such a response (Kenyon and Kriekhaus, (d) impairment i n the a c q u i s i t i o n of a 1-way  1965);  active avoidance response  (Vanderwolf, 1964); (e) a d e f i c i t i n both preoperatively and postoperatively acquired passive avoidance responses (Kaada, Rasmussen, and Kevin, 1962); (f) increased motivation for sucrose or saccharin (Beatty and Schwartzbaum, 1968;  Carey, 1967); (g) decreased  reinforcement  e f f i c i e n c y on a schedule of d i f f e r e n t i a l  at low rates accompanied by an increased rate of  on a f i x e d i n t e r v a l schedule  responding  ( E l l e n and Powell, 1962).  V i r t u a l l y a l l investigators have reported that septal ablation r e s u l t s i n an overt change i n behaviour termed variously as hyperemotionality, septal rage, or h y p e r i r r i t a b i l i t y .  F i r s t observed by Brady and Nauta (1953), i t  consists of an exaggerated s t a r t l e or fear response to previously neutral s t i m u l i and often v i c i o u s attack behaviour i n response to normal handling. Generally, this aspect of the septal syndrome dissipates within a month i f the animals are not handled and are housed separately although the duration may  be reduced to as l i t t l e as two weeks by handling or colony  (Heller, Harvey and Moore 1962;  Brady and Nauta, 1953).  housing  L i n t s , (1965) however,  found that septal lesions produced an increased s e n s i t i v i t y to e l e c t r i c shock at a time when the overt expression of i r r i t a b i l i t y had disappeared.  completely  This i s taken to indicate that septal damage produces long  term a f f e c t i v e changes.  Other i n v e s t i g a t o r s , however, have raised doubts concerning of the s e p t a l area i n the control of emotionality.  the r o l e  McCleary (1961) disputes  the v a l i d i t y of the designation "hyper emotionality" proposed by King (1958), suggesting rather that the apparently a f f e c t i v e changes observed following septal lesions are due to a loss of response i n h i b i t i o n .  This view i s  - 2 supported by Kaada; Rasmussen, and Kevin (1962) who  c i t e as evidence the  contrasting pattern of changes found i n passive versus active shock avoidance behaviour. data suggesting  Kaada, (1960) had e a r l i e r reported e l e c t r o p h y s i o l o g i c a l  that the septal area i s involved i n the control of somatic  responsiveness.  Schwartzbaum, K e l l i c u t and Thompson, (1964) have concluded  that the septal region i s important i n the normal i n h i b i t i o n of reinforced operant responses.  non-  This i n h i b i t i o n i s postulated "to operate  d i r e c t l y upon the expression of highly integrated somatomotor a c t i v i t y . "  Lending support to the septal hyperemotionality  concept are the  r e s u l t s of studies by King and Meyer, (1958) and Schwartzbaum and Gay, These investigators found that amygdaloid lesions appear to reverse exaggerated s t a r t l e response and v i c i o u s reaction to handling lesions.  However, no reversal or change was  caused by septal  situation.  lesions of the amygdaloid n u c l e i are known to r e s u l t i n an (Wood, 1958;  the  observed i n the impairment of  response i n h i b i t i o n s i n a fixed i n t e r v a l reinforcement s h i f t opposite  (1966).  Since  emotionality  i n d i r e c t i o n to that observed following septal ablation  K a r l i , 1956), these studies might be taken as evidence that the  septal syndrome i s not a unitary phenomenon and that "true" emotionality changes co-exist alongside other changes connected to response i n h i b i t i o n impairment.  In f a c t , Carey, (1967) suggests that some of the e f f e c t s of  septal lesions are dependent on locus and there i s evidence (Petsch, Stumpf and Gogolak, 1962;  Donovick, 1968)  of s p e c i f i c nucleus function.  Generally  the reports of wildness and viciousness involve large lesions and  extensive  damage to the l a t e r a l and medial septal n u c l e i but studies by Harrison  and  Lyon (1957) and Thomas, Moore, Harvey and Hunt, (1959) have f a i l e d to i d e n t i f y any precise points involved i n producing the "Emotionality" changes.  The object of the present  i n v e s t i g a t i o n was  to determine whether or  not septal lesions produce a true a f f e c t i v e change i . e . defined p h y s i o l o g i c a l l y as well as behaviourally. hyperemotionality  hyperemotionality  The concept of septal  as employed by both i t s proponents and opponents  i s based almost e x c l u s i v e l y on behavioural  data.  Unfortunately  i t is well  known that frequently employed indices of emotionality such as open f i e l d behaviour, defecation, latency to emerge from a home cage etc. have often f a i l e d to correlate (King, 1968).  In a study designed to determine r e l a t i o n -  - 3 ships between such i n d i c e s w i t h septalectomized animals, Singh, (1966) concludes that "Emotionality i s not unidimensional and d i f f e r e n t i n d i c e s used to assess e m o t i o n a l i t y i n r a t s do not measure the same t h i n g . "  He  found f o r example that whereas s e p t a l r a t s take longer to leave t h e i r home cages, i n d i c a t i n g t i m i d i t y , they are not f e a r f u l i n t h a t they remain i n the open f i e l d f o r longer periods of time. While behavioural measurement of e m o t i o n a l i t y presents the above type of problem, p h y s i o l o g i c a l i n d i c e s are t h e o r e t i c a l l y more r e l i a b l e .  I f we  define e m o t i o n a l i t y as the c o n s t e l l a t i o n of p h y s i o l o g i c a l events t r i g g e r e d by s t r e s s f u l s t i m u l a t i o n , then we can s p e c i f y with c e r t a i n t y what these changes w i l l be.  The general p i t u i t a r y - a d r e n a l response to s t r e s s i s  r e l a t i v e l y w e l l understood and p r e d i c t a b l e (Selye, 1950).  I t would seem  appropriate then, to examine the e f f e c t of s e p t a l l e s i o n s on some aspects of t h i s response.  Kasper, (1965) hypothesized that s e p t a l damage would  r a i s e the l e v e l of p i t u i t a r y - a d r e n a l f u n c t i o n i n response to s t r e s s f u l stimulation.  Her study, which involved s t r e s s i n g s e p t a l r a t s at 5-10  minute  i n t e r v a l s d a i l y v i a a i r b l a s t s f o r one post operative month, produced a number of c o n t r a d i c t i o n s , however.  Heavy p i t u i t a r i e s and adrenals and a  very high p i t u i t a r y ACTH content i n l e s i o n e d unstresses! animals  suggested  p i t u i t a r y - a d r e n a l hyperfunction, while heavy thymuses atad low blood cort i c o s t e r o n e l e v e l s i n s t r e s s e d s e p t a l animals suggested hypo f u n c t i o n under stress.  Kasper suggests that these seeming i n c o n s i s t e n c e s may be due to the  e f f e c t s of a l t e r e d behaviour on the relevant p h y s i o l o g i c a l responses  interacting,  with the d i r e c t e f f e c t s of neural damage. Measurement of d i r e c t manifestations of the pitmriLtary-adrenal f u n c t i o n as u t i l i z e d by the Kasper study i s l i m i t e d by the requirement animal be s a c r i f i c e d i n order to obtain the data.  that the  Consequently  only one  measurement per subject i s p o s s i b l e . With regard to s e p t a l damage, where the behavioural "hyperemotionality" attenuates with tiime i t would be p a r t i c u l a r l y valuable to be able to make repeated meas.ma?es of some physiol o g i c a l index of e m o t i o n a l i t y . One index which lends i t s e l f w e l l to such a procedure i s the hyperglycaemic  response to s t r e s s i.e-„  the r a p i d r i s e i n  blood glucose concentration normally observed i n mammals minutes a f t e r a  - 4 s t r e s s f u l stimulus has been a p p l i e d .  Well documented (Selye, 1950), t h i s  index represents a response d i r e c t l y dependent on changes i n p i t u i t a r y adrenal f u n c t i o n and because only a small blood sample i s required f o r a n a l y s i s , the procedure can be repeated i n d e f i n i t e l y .  Although there i s normally  great v a r i a t i o n i n the r e s t i n g blood glucose l e v e l s of i n d i v i d u a l members of the same species, t h i s source of e r r o r i s minimized by c a l c u l a t i n g a percentage increase due to s t r e s s f o r each animal rather than an absolute increase f o l l o w i n g i t . Experiment I then, involved determining the hyperglycaemic response of s e p t a l r a t s subjected to a period of unavoidable foot shock.  Since the  glucose l e v e l f o l l o w i n g s t r e s s r i s e s to a peak and then declines over time, the i n t e r v a l between onset of foot shock and blood sampling was chosen to provide the maximal post shock glucose concentration (Delay and S o u l i r a c , 1944). period,  The procedure was repeated four times over a s i x week post operative t h i s being the period during which the overt behavioural i r r i t a b i l i t y  i n unhandled animals generally disappears.  Although i n g e s t i o n of large  q u a n t i t i e s of water over a short time can s i g n i f i c e n t l y e f f e c t blood s o l u t e concentrations, the expected r e l a t i v e l y minor increase i n water intake of the s e p t a l animals was monitored as a precaution. I f we take behavioural data concerning rage, a t t a c k and s t a r t l e to be a v a l i d i n d i c a t i o n of "hyperemotionality" i n s e p t a l r a t s , we would expect data concerning the hyperglycaemic response to s t r e s s to lead to s i m i l a r conclusions. Septal subjects should manifest exaggerated r e s t i n g blood glucose l e v e l s and a p a r t i c u l a r l y enhanced glycaemic response to s t r e s s .  We would also  expect t h i s phenomenon to taper o f f over time and normal glucose l e v e l s to r e t u r n w i t h i n approximately 5-6 weeks.  I f , however, the behavioural  m o d i f i c a t i o n s are due to a d i s r u p t i o n of somatomotor i n h i b i t i o n processes rather than hyperemotionality, then no s i g n i f i c a n t a l t e r a t i o n s should be apparent since such d i s r u p t i o n i s presumably not accompanied by a l t e r e d pituitary-adrenal activity. Experiment I a l s o allowed examination of the adrenal gland weights a t the time of s a c r i f i c e .  Kasper (1965) reports adrenal hypertrophy i n s e p t a l  r a t s f o l l o w i n g one month of a f a i r l y severe s t r e s s schedule.  Since the  - 5 duration of s t r e s s involved i n the present study i s i n s i g n i f i c a n t compared to the l a t t e r study, noting the presence or absence of adrenal weight gain would allow us to run a check on Kasper's r e s u l t .  She reported that the  adrenals of non stressed s e p t a l r a t s were not s i g n i f i c a n t l y heavier than those of non stressed c o n t r o l s implying that only septalectomy combined with s t r e s s can produce adrenal hypertrophy.  Of course absence of hyper-  trophy i n s e p t a l animals that have not been severely stressed would tend to detract from the hyperemotionality  theory.  Experiment I Subjects Eighteen experimentally naive, male a l b i n o r a t s of the Wistar s t r a i n served as s u b j e c t s .  These were obtained from the U n i v e r s i t y of B r i t i s h  Columbia Zoology Vivarium and weighed 180-219 gm. at the time of surgery. A l l S_s were i n d i v i d u a l l y housed i n s t a i n l e s s s t e e l wire cages (8 i n . x 10 i n . x 8 i n . ) under continuous i l l u m i n a t i o n , received food and water ad l i b . and . were handled only during surgery and subsequent t e s t i n g .  P r i o r to surgery  the S_s were randomly d i v i d e d i n t o equal experimental and c o n t r o l  groups.  Surgery and H i s t o l o g y Surgery was performed under ether anesthesia w i t h the Ss head held s t a t i o n a r y i n a K r i e g model 51200 s t e r e o t a x i c instrument. maintained throughout  Anesthesia was  the s u r g i c a l procedure by p l a c i n g an ether soaked  s t r i p of s t e r i l e cotton over the S_s snout.  B i l a t e r a l s e p t a l l e s i o n s were  produced i n the experimental group by passing a 2ma.  anodal current (d.c.)  f o r 20 sec. through a s t a i n l e s s s t e e l electrode which was I n s l - x coated except f o r the t i p . The cathode was attached to the Ss t a i l . (deGroot, 1959) were: 1.9 mm. midline and 4.3 mm.  a n t e r i o r to bregma, 0.8 mm.  below the surface .of the cortex.  The coordinates  l a t e r a l to the  In the case of the  c o n t r o l S_s, the e l e c t r o d e was lowered i n the same manner but no current was passed.  A n t i - b i o t i c s u l f a n i l a m i d e powder was a p p l i e d to the wound which  was then closed w i t h three or four sutures.  Immediately  f o l l o w i n g surgery  a l l jSs were i n j e c t e d w i t h 0.1 ml of metrazol stimulant ( p e n t y l e n e t e l r a z o l lOOmg/ml) i n t r a - p e r i t o n e a l l y and 0.2 ml of C r y s t i c i l l i n  (penicillin-procaine  suspension) i n t r a m u s c u l a r l y . A l l S_s were allowed a recovery p e r i o d of 7-9 days i n t h e i r home cages p r i o r to the t e s t i n g schedule. At the conclusion of the experiment  the Ss were s a c r i f i c e d and the  adrenal glands and spleen immediately removed, cleaned of connective t i s s u e and weighed.  The b r a i n was removed and placed i n a f o r m a l i n - s a l i n e s o l u t i o n .  P r i o r to h i s t o l o g y the brains were t r a n s f e r r e d to a 10% e t h y l a l c o h o l s o l u t i o n f o r a few days.  They were then frozen w i t h l i q u i d carbon dioxide and  40u sections cut and s t a i n e d w i t h t h i o n i n .  The l e s i o n s were evaluated by  microscopic examination and the non l e s i o n e d c o n t r o l s e c t i o n s examined f o r p o s s i b l e damage or i n f e c t i o n .  - 7Apparatus A l l S_s were s t r e s s e d i n a wooden compartment 10 i n . long x 6 i n . wide x 12 i n . h i g h , matte b l a c k i n colour and having a transparent l u c i t e hinged cover.  The f l o o r consisted of a g r i d of brass rods each ~  and spaced 1/2 i n . apart.  i n . i n diameter  A 250 a current was passed through the g r i d f o r  3 sec. periods at 10 sec. i n t e r v a l s v i a a shock, source b u i l t by the Psychology Dept. workshop.  D e l i v e r y was timed by a Gerbrands model PT-2A 16 mm. tape  timer. Procedure Each S_was tested four times, the f i r s t t r i a l o c c u r r i n g 8-9 days a f t e r surgery and subsequent t r i a l s a t 16, 28 and 40 days a f t e r surgery. The Ss were deprived of food f o r 3 hours p r i o r t o each t r i a l i n order to e l i m i n a t e the e f f e c t of recent food i n t a k e on blood glucose l e v e l s . A t r i a l consisted of p i c k i n g the S out of i t s home cage with a gloved hand and p l a c i n g i t i n a small wooden r e s t r a i n i n g box, out of which only the t a i l could protrude.  The t a i l was immersed i n hot water f o r approximately 30 s e c ,  removed and t r i e d w i t h paper t o w e l l i n g .  The d i s t a l p o r t i o n o f the t a i l was  then amputated w i t h a s c a l p e l at a p o i n t approximately .75 i n . from the t i p . The flow of blood was stimulated by s t r o k i n g the t a i l from base to t i p between f o r e f i n g e r and thumb while e x e r t i n g moderate pressure.  Blood was c o l l e c t e d  i n a small s l i g h t l y concave glass container-out of which e x a c t l y 0.1 ml was micro-pipetted and t r a n s f e r r e d with washing to a t e s t tube c o n t a i n i n g 3.8 ml. of an i s o t o n i c s o l u t i o n of copper sulphate and sodium sulphate.  The S_ was  then t r a n s f e r r e d to the shock compartment and the shock schedule a c t i v a t e d for  4 minutes.  The S_ then remained i n the compartment f o r a f u r t h e r 5  minutes without r e c e i v i n g shock a f t e r which a second blood sample was taken and the S_ replaced i n i t ' s home cage.  The second blood sampling i n v o l v e d  merely removing the scab formed over the f i r s t c u t . Blood glucose concentrations were determined by a modified v e r s i o n of the Asatoor and King (1954) a l k a l i n e copper reduction method.  To the mixture  of blood and i s o t o n i c copper sulphate-sodium sulphate s o l u t i o n was added 0.1 ml. of 10% sodium tungstate.  This was centrifuged f o r 10 minutes and  1 ml. o f the supernatant removed and added to 1 ml. of a l k a l i n e t a r t r a t e solution.  This was heated i n a b o i l i n g water bath f o r 10 minutes, cooled  - 8 and to i t added 3 ml. of phosphomolybdic s o l u t i o n and 3 ml. of water.  The  r e s u l t i n g s o l u t i o n was mixed and read a f t e r 5 minutes i n a model K l e t t colorimeter using a red f i l t e r .  At the same time 1 ml. of a working  standard of known glucose concentration and a blank using 1 ml. of i s o t o n i c sulphate s o l u t i o n were put up i n the same way. Mg. of glucose per hundred ml. of blood was c a l c u l a t e d as f o l l o w s : e.g. Reading of unknown - reading of blank  ^ ^  n n  Reading of known standard - reading of blank where the known standard i s = 2.5 mg/100 ml. As w e l l as blood sampling, a record of d a i l y water i n t a k e was kept f o r a l l Ss during the f i r s t 3 post-operative weeks. The Ss were s a c r i f i c e d approximately 1 month a f t e r the l a s t t r i a l i  RESULTS - EXPERIMENT I Resting blood glucose The mean r e s t i n g blood glucose concentrations of both s e p t a l e c tomized and c o n t r o l Ss over 4 t r i a l s plus the t o t a l mean f o r each group are summarized i n Table 1. A l l means f a l l w i t h i n what i s considered to be the normal range f o r t h i s measure i n the r a t (60-125 mg. per 100 ml.).  No  s i g n i f i c a n t d i f f e r e n c e s were found between the t o t a l means nor between i n d i v i d u a l groups between or w i t h i n groups f o r any given t r i a l s .  Septal  Ss then do not manifest abnormally high or low r e s t i n g l e v e l s of blood glucose. Change i n blood glucose f o l l o w i n g s t r e s s Corelated _t tests revealed that both s e p t a l and c o n t r o l groups r e g i s t e r e d s i g n i f i c a n t increases i n blood glucose i n response to s t r e s s on a l l t r i a l s .  Table 2 l i s t s the mean absolute increases and t h e i r  signifi-  cance l e v e l s . There was great v a r i a b i l i t y among the blood glucose responses to s t r e s s of i n d i v i d u a l S_s when expressed as percentage changes.  Among the  septalectomized animals t h i s measure showed a range from an increase of 145% to a decrease of 17%; among the c o n t r o l s from an increase of 77% to a decrease of 24%. The t o t a l number of decreases observed was s m a l l however, amounting to f i v e out of seventy-two observations.  I n a l l p r o b a b i l i t y these p a r a d o x i c a l  drops i n the glucose l e v e l do not represent true decreases to response to stress.  I t i s known that i f an animal whose blood sugar i s elevated due to  s t r e s s continues to be s t r e s s e d , r a p i d glucose u t i l i z a t i o n w i l l soon r e s u l t i n a drop to below the pre s t r e s s l e v e l . t h i s occurred i n the present study. recorded.  I t i s l i k e l y t h a t on f i v e occasions  Only two increases of over 100% were  There was l i t t l e consistency on the part of i n d i v i d u a l subjects who  demonstrated very high or negative changes.  For example, one animal recorded  a decrease of 17% on t r i a l 1 and an increase of 61% on t r i a l 3.  On the other  hand another subject recorded an increase of 77% on t r i a l one but an increase of merely 5% on t r i a l 3.  This type of f l u c t u a t i o n occurred among both s e p t a l s  and c o n t r o l s . The mean percent changes i n blood glucose concentration over the 4 t r i a l s as w e l l as the t o t a l mean f o r each group are presented i n Table 3.  - 10 An a n a l y s i s of variance on t h i s data i s summarized i n Table 4. main e f f e c t due e i t h e r to septalectomy or t r i a l s was found.  No s i g n i f i c a n t  However, there -  was a s i g n i f i c a n t i n t e r a c t i o n between septalectomy and t r i a l s .  I n order to  s p e c i f y t h i s i n t e r a c t i o n a f u r t h e r a n a l y s i s of variance f o r simple e f f e c t s was performed and i s summarized i n Table 5.  I t was found that there were  no s i g n i f i c a n t d i f f e r e n c e s between septalectomized and c o n t r o l animals f o r any t r i a l .  Nor were there s i g n i f i c a n t differences between t r i a l s w i t h i n the  s e p t a l or c o n t r o l groups.  For example, T r i a l 1 (Septal-44.3%) does not d i f f e r  from T r i a l 1 (Control-27.7%) or from T r i a l 4 (Septal-20.4%). By v i s u a l i n s p e c t i o n , then, the i n t e r a c t i o n between septalectomy and t r i a l s i n d i c a t e s that during the i n i t i a l  t r i a l s the s e p t a l animals showed  a higher degree of per cent glucose change than the controls although these d i f f e r e n c e s are not s i g n i f i c a n t .  This numerical d i f f e r e n c e a l l but disappears  by T r i a l 4. The great V a r i a b i l i t y i n the data suggests that t h i s s i g n i f i c a n t i n t e r - a c t i o n may represent a Type I e r r o r . Adrenal weights and water intake Table 6 summarizes the mean adrenal weights at the conclusion of the experiment and the mean d a i l y water intake during the study.  Both  measures are r e l a t e d to t o t a l body weight (determined at the time of sacrifice).  As expected, the septalectomized animals drank s i g n i f i c a n t l y  more than the c o n t r o l subjects (p  .05). However, casual observation i n d i c a t e d  that the s e p t a l animals d i d not ingest abnormally large amounts of water over short periods of time.  As mentioned e a r l i e r , i t i s reasonably safe to  assume that the blood glucose l e v e l s of the septalectomized group were not a f f e c t e d by higher water intake l e v e l s . No s i g n i f i c a n t d i f f e r e n c e between the adrenal gland weights of the s e p t a l and c o n t r o l animals was apparent.  - 11 Table 1 Mean r e s t i n g blood glucose c o n c e n t r a t i o n , i n mg. per 100 ml. Trial 3  Trial 4  Trial 1  Trial 2  Septal  63.9  81.4  98.0  85.2  82.1  Control  63.1  89.4  102.1  91.3  86.4  Total  Table 2 Mean r i s e i n blood glucose concentration i n mg. per 100 ml. f o l l o w i n g e l e c t r i c shock  Septal _t  P_  Trial 1  Trial 2  18.5  29.3  30.7  16.0  2.368  5.1974  4.2831  4.4236  <.01  4.02  <-.01  Trial 2  Trial 3  Trial 4  .05  Trial 1 Control _t  Trial 3  Trial 4  16.3 .  21.4  14.8  16.0  2.7084  3.3642  3.0382  3.2916  -e .05  €.01  «".02  €.02  Table 3 Mean % change i n blood glucose concentration f o l l o w i n g e l e c t r i c shock Trial 1  Trial 2  Trial 3  Trial 4  Total  Septal  44.3  39.8  36.0  20.4  35.1  Control  27.7  25.0  15.1  18.7  21.6  - 12 -  Table 4 A n a l y s i s of variance (two way with r e p l i c a t i o n on one f a c t o r ) % change i n Blood glucose concentration Source  df  Between Gp.  17  24006.1  1411.7  1  3307.6  3307.6  E r r o r between  16  20698.5  1293.7  Within Gp.  .54  34003.5  629.7  Trials(T)  3  2864.6  954.8  1.91  T x S  3  7091.7  2363.9  4.72  24047.2  501,0  Septalectomy(S)  Error within  48  ss  ms  I  2.  2.56  t-01  - 13-  Table 5  A n a l y s i s of variance f o r simple e f f e c t s % change i n blood glucose concentration df  ss  ms  At t r i a l 1  1  34.4  34.4  At t r i a l 2  1  27.4  27.4  At t r i a l 3  1  54.6  54.6  At t r i a l 4  1  0.3  0.3  Within c e l l  64  Source Between Gp.  20698.5  323.4  Within Gp. Between t r i a l s for s e p t a l s  80.8  26.9  Between t r i a l s for c o n t r o l s  24.9  8.3  Septalectomy x trials Trials x subjects w i t h i n Gp.  48  7091.7  2363.9  26806.1  558.5  - 14 -  Table 6 Mean adrenal weights and mean d a i l y water Intake corrected f o r body weight. Adrenal Wt. mgm. per gm. Septal Control *  Water intake ml. per gm.  ' 1.074  0.1707 *  0.812  0.1245 *  t=2.5054 df=15 p-^.05 (two t a i l e d t e s t )  Experiment I I Experiment I I digresses somewhat from a d i r e c t p h y s i o l o g i c a l approach to the s e p t a l syndrome but s t i l l deals w i t h the nature of changes i n "emotional" behaviour due to s e p t a l damage.  More p r e c i s e l y , i t would seem of i n t e r e s t to  manipulate some of the behavioural r a t i n g s of e m o t i o n a l i t y (mentioned e a r l i e r ) v i a some agent whose behavioural and p h y s i o l o g i c a l e f f e c t s are r e l a t i v e l y w e l l known.  Rather than concern ourselves with the presence or absence of a  s t a t i s t i c a l c o r r e l a t i o n between such r a t i n g s we intend to focus a t t e n t i o n on how i n d i v i d u a l i n d i c e s are a f f e c t e d by t h i s agent.  Open f i e l d behaviour,  r e s i s t a n c e to handling, latency to emerge from a home cage e t c . would be observed under the i n f l u e n c e of a drug known to i n f l u e n c e such behaviour and thought to have some connection with the l i m b i c system. chlorpromazine hydrochloride.  Such a drug i s  A depressant, i t diminishes spontaneous  motor a c t i v i t y (Goodman and G i l l a m , 1957) but does not n e c e s s a r i l y impair behaviour; e.g. where m o b i l i t y i s desireable animals treated with c h l o r promazine may perform b e t t e r than c o n t r o l s (Blough, 1958).  Bindra and Bacon,  (1959) found that chlorpromazine decreased some components of general a c t i v i t y while l e a v i n g others unaffected. Conditioned avoidance responses are blocked by i t although the unconditioned response to the noxious stimulus remains unaffected (Cook and Weidley, 1957).  Nelson and F r a n k l i n , (1967) found that  chlorpromazine f a c i l i t a t e d fear e x t i n c t i o n w h i l e Schallek, Kuehn and Jew (1962) reported depressed viciousness i n s e p t a l r a t s upon a d m i n i s t r a t i o n of a v a r i e t y of compounds s i m i l a r chemically to chlorpromazine. K i l l a m and K i l l a m , (1958) found that chlorpromazine may depress the arousal response of the l i m b i c system that i s c h a r a c t e r i z e d by slow high voltage waves.  I t may a l s o slow spontaneous l i m b i c a c t i v i t y (Schallek,  1962)  and has an i n h i b i t o r y e f f e c t on s e p t a l s e l f s t i m u l a t i o n i n doses that do not disorganize behaviour generally (Olds and T r a v i s , (1960); Olds,,I960).  In  r a t s having s e p t a l area l e s i o n s there i s u s u a l l y an increase i n low amplitude c o r t i c a l EEG a c t i v i t y and a degrease i n the number of d.c. s h i f t s which can be blocked by the a d m i n i s t r a t i o n of chlorpromazine ( P i r c h and Norton, 1967). C l i n i c a l l y phlorpromazine i s widely employed to reduce anxiety and has been found to reduce p a i n s e n s i t i v i t y ( N i c k o l s , 1960).  Kletzkin,  (1962)  suggests that the therapeutic value of chlorpromazine i s due to i t s d i s r u p t i o n  - 16 of  abnormal n e u r a l p a t t e r n s i n the l i m b i c  Experiment lesions  I I then i s designed  of  septal  of  chlorpromazine administration  system.  to r e p l i c a t e once a g a i n the  on " e m o t i o n a l " b e h a v i o u r on s u c h  and  to i n v e s t i g a t e  behaviour.  the  effects  effects  METHOD Subjects The Ss were 16 experimentally naive male a l b i n o r a t s weighing 350380 gm. a t the time of surgery.  Other than the f a c t that they were obtained  from Simonsen Breeding Laboratories (Los Angeles) a l l d e t a i l s were as i n Experiment I . Surgery and H i s t o l o g y . S u r g i c a l and h i s t o l o g i c a l procedures were i d e n t i c a l to those employed i n Experiment I except that Nembutal (sodium p e n t o b a r b i t a l ) anesthesia (1.5 mg./ 100 gm.) was u t i l i z e d i n place of ether and that the adrenal glands were not examined. Apparatus The apparatus consisted of an open f i e l d with i n t e r n a l dimensions 4 x 4 x 1 f t . The f l o o r and w a l l s were painted with white enamel and the f l o o r surface marked o f f i n t o s i x t e e n equal squares by black l i n e s , 1/8 i n . i n width.  approximately  A black, matte wooden box (6 x 10 x 12 i n . ) w i t h a hinged  top was attached to one side of the open f i e l d and was separated from i t by a 3 x 4 i n . black g u i l l o t i n e door.  The e n t i r e apparatus was s i t u a t e d i n a small  room having a matte black c e i l i n g and matte dark grey w a l l s .  I l l u m i n a t i o n was  provided by a 15 watt bulb held i n a black paper c y l i n d e r suspended 5 f t . above the open f i e l d . Procedure: Each S_ was given two t r i a l s , one f o l l o w i n g a d m i n i s t r a t i o n of c h l o r promazine, the other f o l l o w i n g a d m i n i s t r a t i o n of s a l i n e .  The f i r s t t r i a l  occurred 10-12 days a f t e r surgery, the second 2 days f o l l o w i n g T r i a l 1. The order of t e s t i n g was counter balanced so that h a l f of each group received chlorpromazine  f i r s t and s a l i n e second and the other h a l f s a l i n e f i r s t and  chlorpromazine  second.  A t r i a l consisted of s l i d i n g S_s cage out of the  colony rack, covering the open top and c a r r y i n g i t i n t o the t e s t room where i t was l e f t undisturbed f o r 10 minutes i n a p o s i t i o n that d i d not allow the S_ to see the apparatus.  The S_was then picked up w i t h a gloved hand and  i n j e c t e d i n t r a p e r i t o n e a l l y w i t h chlorpromazine  (.2 mg/lOOgm i n 1.6 ml d i s t i l l e d  water) or a l i k e amount of s a l i n e and replaced i n the cage f o r 20 minutes. At the end of t h i s period the S_ was picked up and was rated on a 1-7 scale f o r  - 18 r e s i s t a n c e to capture and on the same s c a l e f o r r e s i s t a n c e t o handling f o l l o w i n g capture.  Resistance was considered i n terms of locomotion,  v o c a l i z a t i o n , b i t i n g , clawing and s t r u g g l i n g .  The S_ was held f o r not more  than approximately 20 sec. and then placed i n t o the home box which was closed completely f o r 2 minutes.  The g u i l l o t i n e door was then removed and  the number of times S_s head emerged from the box and the number of times S_ placed approximately h a l f i t s body i n c l u d i n g both f o r e l e g s over the threshold were observed during a 10 min. p e r i o d . At the end of t h i s session the g u i l l o t i n e door was replaced, S_ removed through the top of the home box and placed back i n i t s cage.  Two minutes were allowed to elapse a f t e r  which S_ was picked up and placed onto one corner of the open f i e l d , the corner.  facing  During the next 5 minutes S_ was observed from one side and the  number of squares traversed, the number of hind l e g rears and the number of f e c a l boluses dropped were noted.  At the conclusion of t h i s s e s s i o n S_ was  replaced i n i t s cage and returned to the animal colony.  Between t r i a l s ,  the open f i e l d was washed with a s o l u t i o n of water and vinegar i n order to e l i m i n a t e scent t r a i l s .  RESULTS Experiment I I Resistance to capture and handling Table 7 provides a summary of mean r a t i n g s of r e s i s t a n c e to capture and handling on 1 - 14 s c a l e .  At the time of observation separate r a t i n g s  for r e s i s t a n c e to capture and r e s i s t a n c e to handling were made on a 1 - 7 scale.  However, since i t i s d i f f i c u l t to s p e c i f y when capture becomes  handling, i t seemed prudent to combine the capture and handling scales f o r each animal.  "Resistance" i s a somewhat s u b j e c t i v e term.  Factors taken  i n t o c o n s i d e r a t i o n i n c l u d e as i n d i c a t e d e a r l i e r ; v o c a l i z a t i o n , u r i n a t i o n and defecation, locomotion (escape), squirming, b i t i n g and s c r a t c h i n g . A given animal may, f o r example, have manifested only squirming or v o c a l i z a t i o n and yet r a t e a high score on the b a s i s of i n t e n s i t y . The table i s arranged so that i t i s evident how the order of a d m i n i s t r a t i o n of chlorpromazine and s a l i n e a f f e c t e d the group means.  This  form of t a b u l a t i o n i s u t i l i z e d f o r the other measures i n Experiment I I as well.  A L a t i n Square a n a l y s i s of variance (Bruning and K i n t z , 1967) was per-  formed on t h i s data (again t h i s type of a n a l y s i s i s u t i l i z e d Experiment  I I ) and i s summarized i n Table 8.  throughout  Septalectomized subjects were  found to r e s i s t capture and handling to a greater extent than c o n t r o l animals (p x .05). group.  Chlorpromazine a d m i n i s t r a t i o n , however, had no e f f e c t on e i t h e r  Neither d i d an order e f f e c t appear, there being no s i g n i f i c a n t  d i f f e r e n c e i n the degree of r e s i s t a n c e on f i r s t and second t r i a l s i n e i t h e r group.  No s i g n i f i c a n t i n t e r a c t i o n s occurred.  Open f i e l d behaviour Squares traversed The mean number of squares traversed i n an open f i e l d apparatus over 5 minutes are i n d i c a t e d i n Table 9 w h i l e Table 10 provides a summary of the a n a l y s i s of variance on the data.  Septalectomized Ss traversed f a r fewer  squares than the operated c o n t r o l s and t h i s d i f f e r e n c e was h i g h l y s i g n i f i c a n t (p i: .001).  Within groups order e f f e c t was found as w e l l , i n d i c a t i n g that  among both s e p t a l and c o n t r o l subjects the f i r s t t r i a l produced a higher score than the second t r i a l regardless of whether s a l i n e or chlorpromazine was  - 20 utilized.  Apparently, the i n i t i a l 5 minute exposure to the open f i e l d plus  the 10 minute opportunity to observe the apparatus from the home box r e s u l t i n a degree of h a b i t u a t i o n that a f f e c t s the s e c o n d . t r i a l . The a d m i n i s t r a t i o n of chlorpromazine d i d not a f f e c t the scores of e i t h e r group, however, a s i g n i f i c a n t between groups drug by order i n t e r a c t i o n did occur.  This means that the d i f f e r e n c e between r e c e i v i n g chlorpromazine  f i r s t and chlorpromazine second i s greater among the operated controls than among the s e p t a l s u b j e c t s . Rears Aside from simple ambulation, r e a r i n g up on the handlegs i s i n the rat  the only other d i s t i n c t motor p a t t e r n that we may observe as c o n t r i b u t i n g  to exploratory or general a c t i v i t y i n an open f i e l d .  I t i s not s u r p r i s i n g  then that r e s u l t s with regard to the number of rears recorded should p a r a l l e l the "squares t r a v e r s e d " data.  Tables 11 and 12 summarize the number of rears  i n the open f i e l d 5 minute s e s s i o n and provide the r e s u l t s of the a n a l y s i s of variance.  I t was found that septalectomized Ss demonstrated  fewer rears  (p '<. .025) then the operated controls and that i n both these groups higher scores were recorded on the f i r s t t r i a l than on the second.  Again no e f f e c t  i s a t t r i b u t a b l e to the a d m i n i s t r a t i o n of chlorpromazine and u n l i k e the "squares t r a v e r s e d " scores no s i g n i f i c a n t i n t e r a c t i o n s occurred. j Defecation Tables 13 and 14 summarize the mean number of boluses dropped during the 5 minute open f i e l d s e s s i o n and the mean number of times the S_'s head emerged from a home box opening on the open f i e l d , r e s p e c t i v e l y .  A n a l y s i s of  variance revealed no s i g n i f i c a n t e f f e c t s or i n t e r a c t i o n s . Half e x i t s The mean number of times i n 10 minutes that the j>s performed a h a l f e x i t by p l a c i n g both forelegs over the threshold of the home box are l i s t e d i n Table 15. A l l the operated c o n t r o l s and a l l the septalectomized Ss that were run under the chlorpromazine treatment had zero scores.  Only the septalectomized  S_s run under the s a l i n e c o n d i t i o n r e g i s t e r e d p o s i t i v e scores.  I t would appear  - 21 -  then that s e p t a l r a t s have a tendency t o leave present i n normal animals. course, of  Chlorpromazine suppresses t h i s  tendency.  t h e s m a l l number o f h a l f e x i t s a c t u a l l y r e c o r d e d makes t h i s  conclusion  highly speculative.  T h e same r e s e r v a t i o n s h o u l d  to the f a c t that of the s e p t a l animals ones t h a t that  a home c a g e w h i c h i s n o t  received  received  2.  kind apply  that d i d r e g i s t e r p o s i t i v e scores,  t h e s a l i n e i n j e c t i o n on t r i a l  i t on t r i a l  also  Of  1 scored  higher  the  than those .  - 22 Table 7. Mean r a t i n g of r e s i s t a n c e to capture and handling on a 1-14 s c a l e CPZ -chlorpromazine hydrochloride SAL - i s o t o n i c s a l i n e Septals order of admin-istration  CPZ  SAL  s i - s4  CPZ - SAL  9.5  5.8  s5 - s8  SAL - CPZ  9.5  12.0  9.5  8.9  order of admin-istration  CPZ  SAL  s9 - s l 2  CPZ - SAL  4.5  5.0  sl3 - sl6  SAL - CPZ  6.3  7.0  5.4  6.0  s i - s8  Controls  s9 - s l 6  - 23 Table 8 A n a l y s i s of variance ( L a t i n Square) Resistance to handling and capture Source  df  Between gp.  15  Sum of squares  Mean squares  Septalectomy (S)  1  98.0  98.0  5.2  CPZ x Order ( 0 )  1  50.0  50.0  2.67  CPZ x S  1  3.15  3.15  Error  12  224.7  18.73  Within Gp.  16  130.0  CPZ  1  Order  1  21.2 '  CPZ x S  1  3.15  0 x S  1  18.0  Error  12  87.7  0  0 21,2 3.15 18.0 7.31  0.17  0 2.9 0.43 '2.46  ^ .05  - 24 Table 9 Mean number of squares crossed i i i an open f i e l d apparatus during a f i v e min. p e r i o d . CPZ - chlorpromazine hydrochloride SAL - i s o t o n i c s a l i n e Septals Order of administration  CPZ  SAL  0  si  s4  CPZ - SAL  12.5  s5  s8  SAL - CPZ  4.5  5.8  si  s8  8.0  2.9  Controls Order of adminis t r a t i o n  CPZ  SAL  s9 - s l 2  CPZ - SAL  56.5  47.3  sl3 - sl6  SAL - CPZ  8.5  35.8  32.5  41.5  s9 - s l 6  - 25 Table 10 -Analysis of variance ( L a t i n Square) NO. of squares crossed i n an open f i e l d apparatus during a 5 min. period. Source  df  Siim of squares  Mean squares  F  Between Gp.  15  16119.9  -  -  Septalectomy (S)  1  7969.5  7969.5  20.75  *.001  CPZ x Order (0)  1  1845.2  1845.2  4.8  «-.05  CPZ x S  1  1696.6  1696.6  4.42  Error  12  4608.6  384.1  -  Within Gp.  16  4321.5  240.1  -  CPZ  1  30.0  30.0  0.15  Order  1  1212.7  1212.7  6.07  CPZ x S  1  399.0  399.0  1.99  0 x S  1  282.0  282.0  1.41  Error  12  2397.6  199.8  -  P_  <:.05  - 26 Table 11 Mean number of- rears i n an open f i e l d apparatus during a 5 min. period. CPZ - chlorpromazine hydrocholride SAL - i s o t o n i c s a l i n e Septals Order of adminis t r a t i o n  CPZ  SAL  s i - s4  CPZ - SAL  5.3  0  s5 - s8  SAL - CPZ  5.5  5.8  5.4  2.9  CPZ  SAL  s i - s8 Controls Order of administration s9 - s l 2  CPZ - SAL  21.0  11.0  sl3 - sl6  SAL - CPZ  6.0  14.7  13.5  12.9  s9 - s l 6  Table 12 ..' A n a l y s i s of Variance (Latin. Square) No. of rears i n an open f i e l d apparatus during a 5 min. period. Source  df  Sum of squares  Mean •squares  • p  Between Gp.  15  1422.7  Septalectomy (S)  : l  657.0  657.7  8.17  CPZ x Order (0)  l  13.8  13.8  0.17  CPZ x S  l  148.0  148.0  1.84  Error  12  965.2  80.4  -  Within Gp.  16  634.5  -  -  CPZ  1  19.5  19.5  1.03  Order  1  294.0  294.0  CPZ x.S  1  7.1  7.1  0.38  0 x S  1  87.8  87.8  4.6  Error  12  226.1  18.8  —  -  15.6  - 28 Table 13 Mean number of boluses excreted i i i an open f i e l d apparatus during a f i v e minute period. CPZ 2. chlorpromazine hydrochloride SAL - i s o t o n i c s a l i n e Septals Order of administration  CPZ .  SAL  s i - s4  CPZ - SAL  1.25  0.5  s5 - s8  SAL - CPZ  0.5  1.0  0.88  0.75  CPZ  SAL  s i - s8 Controls Order of administration s9 - s l 2  CPZ - SAL  1.0  0.25.  sl3 - sl6  SAL - CPZ  0.75  1.25  0.88  0.75  s9 - s l 6  '  29 .Table 14 Mean number of head emergences from a home box during a 10 min. p e r i o d . CPZ - chlorpromazine hydrochloride SAL - i s o t o n i c s a l i n e Septals Order of administration  CPZ  SAL  s4  CPZ - SAL  1.5  3.5  s5 - s8  SAL - CPZ  2.5  9.8  2.0  5.7  CPZ  SAL  •si -  s i - s8 Controls Order of administration s9 - s l 2  CPZ - SAL  3.0  2.0  sl3 - sl6  SAL - CPZ  2.0  2.0  2.5  2.0  s9 - s16  - 30 Table 15 Mean number of times both f o r e l e g s were placed over the threshold of a home box e x i t during a 10 miri. p e r i o d . Septals Order of administration  CPZ  SAL  s i - s4  CPZ - SAL  0  1.5  s5 - s8  SAL - CPZ  0  5.0  0  3.3  CPZ  SAL  s i - s8 Controls Order of adminis t r a t i o n s9 - s l 2  CPZ- - SAL  0  0  sl3 - sl6  SAL - CPZ  0  0  0  0  s9 - s l 6  - 31 -  Anatomical f i n d i n g s - Experiments I and I I Appendix I contains h i s t o l o g i c a l reconstructions (deGroot, of s e p t a l damage i n each lesioned subject. per b r a i n are shown 6.2, tively.  7.0,  7.8,  1959)  Four representative sections  and 8.6 mm.  a n t e r i o r to bregma respec-  Examination shows that a l l l e s i o n s caused extensive damage to the  l a t e r a l and medial s e p t a l n u c l e i . to the f o r n i x .  S i x of the l e s i o n s included some damage  In c e r t a i n cases there appeared to be v i a b l e s t r i p s of  t i s s u e between the l a t e r a l n u c l e i , however, these do not appear i n the reconstructions since they were l o s t i n the s e c t i o n i n g process. there was  In general  a high degree of s i m i l a r i t y i n the extent and shape of the l e s i o n s .  The brains of Ss from the c o n t r o l group were sectioned and examined f o r signs of i n f e c t i o n or other damage, however, none were found.  Discussion and Conclusions Assuming that a r i s e i n the concentration of blood glucose i s a v a l i d i n d i c a n t of a change In a r a t ' s emotional s t a t e , the absence of any changes i n e i t h e r the r e s t i n g glucose l e v e l or the hyperglycaemic response to s t r e s s f o l l o w i n g s e p t a l l e s i o n s may be construed as evidence that such damage does not lead to hyperemotionality.  The extreme i r r i t a b i l i t y generally reported  i n the wake of l e s i o n s of the septum and confirmed i n Experiment I I must be explained otherwise.  Since the response i n h i b i t i o n theory of the s e p t a l  syndrome i s c u r r e n t l y the only a l t e r n a t i v e hypothesis, i t i s i n d i r e c t l y supported. In defence of the concept of s e p t a l hyperemotionality one might suggest that i t i s unreasonable to expect to observe exaggerated glucose l e v e l s i n s e p t a l animals.  The release of a d d i t i o n a l glucose i n t o the blood stream  during s t r e s s f u l periods would serve n o . r e a l adaptive f u n c t i o n over and above the normal hyperglycaemic response, since the s t r e s s f u l s t i m u l a t i o n presented to the s e p t a l animal i s no more p h y s i c a l l y harmful nor d i f f i c u l t to escape than i t i s to a normal one.  Likewise a supra normal r e s t i n g glucose l e v e l i n  an undisturbed s e p t a l animal would serve no p a r t i c u a l r purpose. Perhaps what should be i n v e s t i g a t e d w i t h regard to blood l e v e l s i n septalectomized subjects i s t h e i r degree of s e n s i t i v i t y to s t i m u l a t i o n .  Can  the hyperglycaemic response be t r i g g e r e d by a lower l e v e l of s t r e s s f u l s t i m u l a t i o n i n a s e p t a l animal than i n a normal one?  In other words, instead of  looking f o r an exaggerated hyperglycaemic response to severe s t i m u l a t i o n we would look f o r a simple hyperglycaemic response to a stimulus which would under ordinary circumstances not warrant such a response.  Should t h i s s i t u a -  t i o n prove to be the case, i t would f a l l conveniently i n t o l i n e w i t h b a s i c behavioural observations. Exaggerated s t a r t l e and attack behaviour, f o r example, which would be expected i n response to intense s t i m u l a t i o n such as a sudden loud noise or a d i r e c t a t t a c k by another animal i s e l i c i t e d by simply opening a cage of a septalectomized animal. The trouble w i t h a threshold approach such as t h i s i s that i t tends to be a restatement of the response i n h i b i t i o n hypothesis modified to include  - 33 -  p h y s i o l o g i c a l responses as w e l l as behavioural ones.. I t places the emphasis on the magnitude of the stimulus required to e l i c i t a given response rather than on the magnitude of the response to a given stimulus.  When dealing w i t h  p h y s i o l o g i c a l i n d i c e s of emotionality or response to s t r e s s , however, i t i s the magnitude of the index rather than i t s threshold which has most o f t e n been e s t a b l i s h e d as the c r i t i c a l f a c t o r (Selye, 1959;  L e v i , 1965).  For example,  l e s i o n s of the amygdaloid area r e s u l t i n increased aggressiveness (Wood or increased p l a c i d i t y (King, 1959)  1958)  depending on the locus of the damage.  Accompanying the former, Bovard and Gloor (1961) and Anand (1957) reported an increase i n the c o r t i c o s t e r o n e response to i m m o b i l i z a t i o n s t r e s s whereas Mason and Nauta (1959) found that the l a t t e r l e s i o n r e s u l t e d i n the v i r t u a l disappearance of the 17-hydroxy c o r t i c o s t e r o i d response to avoidance t r a i n i n g . P o i r i e r , Cordeau, Lemire and Ayotte, (1956) found that b i l a t e r a l amygdaloid damage of the type producing p l a c i d i t y , are followed by a b o l i t i o n of the hyperglycaemic response to s t r e s s .  E l e c t r i c a l s t i m u l a t i o n i n t h i s area  causes an increase i n 17- h y d r o x y - c o r t i c o s t e r o i d production i n monkeys  (Mason,  1958)  ,  Where neural damage i s not i n v o l v e d , the same type of q u a n t i t a t i v e changes have been reported.  Henderson (1967) f o r example, found that r a t s w i t h  high behavioural emotionality scores due to preweaning noxious s t i m u l a t i o n a l s o have high r e s t i n g blood sugar l e v e l s . changes i n endocrine and metabolic  In general, reports of q u a n t i t a t i v e  a c t i v i t y associated with changes i n  e m o t i o n a l i t y or the response to s t r e s s f u l s t i m u l a t i o n are very numerous.  It  i s reasonable to expect, then, that i f s e p t a l l e s i o n s were -indeed producing an increase i n emotionality, t h i s s i t u a t i o n should manifest i t s e l f i n a q u a n t i t a t i v e l y measurable p h y s i o l o g i c a l manner.  As f a r as blood glucose  l e v e l s or adrenal weights are concerned, t h i s has not been the case i n Experiment I. Rasper's (1965) i n v e s t i g a t i o n does, i n f a c t , encompass such f i n d i n g s . However, conclusions drawn from the various measures that were u t i l i z e d i n that study, are c o n t r a d i c t o r y .  C e r t a i n i n d i c e s of p i t u i t a r y adrenal a c t i v i t y  i n d i c a t e d hyperfunction i n s e p t a l animals, others no change at a l l .  At the  same time i t appeared that septalectomy reduces responsiveness to s t r e s s  - 34 although there was s i g n i f i c a n t adrenal hypertrophy.  This l a s t p o i n t d i r e c t l y  c o n t r a d i c t s the f i n d i n g s of the present experiment which revealed no increase i n adrenal weights.  A very probable explanation,, however, i s the simple  d i f f e r e n c e i n the. d u r a t i o n of s t r e s s between.the two s t u d i e s .  Experiment  I, although u t i l i z i n g a f a i r l y severe form of s t r e s s , d i d not expose the subjects to i t f o r anywhere near the l e n g t h of time used by Kasper.  I t may  be concluded that septalectomy i t s e l f does not lead.to gains i n adrenal weight. Kasper, i n f a c t , d i d not f i n d a s i g n i f i c a n t gain i n adrenal weight among her non-stressed s e p t a l animals.  This was confirmed as w e l l by Clody and C a r l t o n  (1968).  •  _  Unfortunately, other i n c o n s i s t e n c i e s w i t h i n the Kasper  ' ' experiments  and between that study and the present one do not y i e l d to c l a r i f i c a t i o n easily.  One p o s s i b i l i t y that bears c o n s i d e r a t i o n i s that the behavioural  m o d i f i c a t i o n s f o l l o w i n g s e p t a l l e s i o n s produce endocrine changes not d i r e c t l y r e l a t e d to the accompanying n e u r a l damage.  As w e l l , the Kasper study r e l i e d  on s i n g l e measures of cumulative e f f e c t s f o l l o w i n g a long p e r i o d of s t r e s s while the present i n v e s t i g a t i o n i n v o l v e d repeated measures of a d i f f e r e n t short term e f f e c t .  I t i s d i f f i c u l t to say which approach i s more v a l i d .  Repeated measures of a temporary change such as the hyperglycaemic  response  seem to be j u s t i f i a b l e because of the t r a n s i e n t nature of the behavioural s e p t a l syndrome. i s i n order.  Perhaps a more d i r e c t index of p i t u i t a r y - a d r e n a l f u n c t i o n  Monitoring the l e v e l s of u r i n a r y k e t o s t e r o i d s f o l l o w i n g  septalectomy i s one p o s s i b i l i t y .  Another might i n v o l v e repeated measures of  plasms c o r t i c o s t e r o n e s i n amounts of blood not required s a c r i f i c e of the subject. Experiment I I generally f a i l e d to shed new l i g h t on the nature of the s e p t a l syndrome.  A d m i n i s t r a t i o n of chlorpromazine a f f e c t e d n e i t h e r s e p t a l  nor operated c o n t r o l animals with respect to r e s i s t a n c e to capture and handling, open f i e l d behaviour or d e f e c a t i o n . Only i n the case of h a l f - e x i t s d i d chlorpromazine appear to suppress the tendency of a s e p t a l animal to emerge from a home box.  Even t h i s e f f e c t i s a dubious one, however, since the number  of h a l f - e x i t s i n v o l v e d was s m a l l .  As w e l l , had f u l l r a t h e r than h a l f e x i t s  been designated as the s c o r i n g c r i t e r i o n no a f f e c t whatsoever would appear  -  - 35 since no animal a c t u a l l y accomplished a f u l l e x i t . Inadequate dosage immediately comes to mind by way of e x p l a n a t i o n . At 0.2 mg. per 100 gm. of body weight, however, the dose u t i l i z e d i n Experiment I I equals or exceeds the amounts administered by most behavioural and physiol o g i c a l i n v e s t i g a t o r s (Cornetsky and Bain, 1965; P i r c h and Norton; 1966; Archer, 1954; Boyd and M i l l e r , 1954). 100 gm  Watson and Steinberg (1967) report that .2 mg/  u s u a l l y produces a very s l i g h t muscular f l a c i d i t y i n r a t s .  A larger  dose then may i n v o l v e r i s k i n g a degree of muscular impairment that would negate other f i n d i n g s .  I t i s i n t e r e s t i n g to note that i n the same study Watson  and Steinberg found that t h i s dosage of chlorpromazine f a i l s to i n h i b i t the hyperglycaemic response to s t r e s s . I t has been reported that chlorpromazine depresses general a c t i v i t y as measured i n a c t i v i t y cages (Kinnard and Carr, 1957) or wheels (Routtenberg and Kuznesoff, 1967) and that i t a f f e c t s s e l e c t e d components of a c t i v i t y (e.g.  s n i f f i n g ) while not a f f e c t i n g others (e.g. grooming) (Bindra and Bacon,  1959).  Apparently, however, i f a more stimulus oriented form of a c t i v i t y  r e l a t e d to the degree of f e a r f u l n e s s or aggressiveness i s examined as was i n the present study, chlorpromazine has no e f f e c t on e i t h e r normal or s e p t a l animals.  P o s s i b l y any depressant or other e f f e c t s of the drug are masked  by the l e v e l of arousal i n the subjects due to the experimental s i t u a t i o n . The e f f e c t s of the l e s i o n s proper were v a r i e d and not n e c e s s a r i l y consistent with p r e v i o u s l y reported f i n d i n g s .  As f i r s t observed by Brady and  Nauta (1953) and o f t e n t h e r e a f t e r , the septalectomized r a t s displayed very high r e s i s t a n c e to capture and handling i n a d e f i n i t e demonstration of s e p t a l rage.  The same animals c l e a r l y manifested l e s s exploratory behaviour than  c o n t r o l s i n an open f i e l d s i t u a t i o n .  However, N i e l s o n , Mclver and Boswell  (1965), Clody and C a r l t o n (1968) and Douglas and Raphelson (1966) have a l l reported increased e x p l o r a t o r y behaviour i n s e p t a l r a t s u t i l i z i n g measures other than open f i e l d behaviour. between s m a l l enclosed spaces.  Their s t u d i e s involved movement w i t h i n or Singh (1966), on the other hand, has found  that i n a s i t u a t i o n where r a t s may  leave an open f i e l d and r e t u r n to a home  - 36 -  cage, s e p t a l animals tend to remain i n the open f i e l d although they do not explore as much as c o n t r o l s .  The N i e l s o n e t a l . study a l s o revealed that  lesioned animals show very l i t t l e a c t i v i t y i n an open f i e l d apparatus. I f we consider a l l of the preceeding data i n the l i g h t of present f i n d i n g s , i . e . that s e p t a l r a t s .... a) are more r e s i s t a n t to handling and capture b) traverse fewer squares i n an open f i e l d c) demonstrate fewer rears i n an open f i e l d d) may have a greater tendency to leave a home cage .... the f o l l o w i n g p i c t u r e emerges.  Septalectomized r a t s , u n l i k e normal  animals, apparently tend to attempt to leave small enclosed spaces, however, when placed i n a b r i g h t open space they are l e s s a c t i v e than c o n t r o l s . How t h i s s i t u a t i o n can be i n t e r p r e t e d i n terms of e m o t i o n a l i t y i s another matter.  Low l e v e l s of exploratory a c t i v i t y i n an open  field  apparatus may be taken as an i n d i c a t i o n of t i m i d i t y or f e a r f u l n e s s ( V a l l e , 1970).  Another approach, however, would view i t as i n d i c a t i v e of a cautious,  aggressive animal.  The concept of reduced response i n h i b i t i o n due to s e p t a l  damage would not seem a p p l i c a b l e here since absence of response rather than i n h i b i t i o n seems to be the key f a c t o r .  facilitation  On the other hand, t h i s  concept would e x p l a i n the tendency to be a c t i v e i n or to leave s m a l l enclosed spaces.  We have o f t e n observed s e p t a l r a t s to leap out of open colony cages,  something r a r e l y seen i n normal r a t s .  This l a t t e r suggestion i s h i g h l y  s p e c u l a t i v e of course, and serves mainly to i l l u s t r a t e the d i f f i c u l t y i n i n t e r p r e t i n g animal behaviour i n e s s e n t i a l l y s u b j e c t i v e human terms such as f e a r f u l and f e a r l e s s . Experiment  I I , then, does not allow f o r s p e c i f i c conclusions. The  f a c t that behavioural i n d i c e s of e m o t i o n a l i t y do not c o r e l a t e w e l l w i t h each other i s borne out to a c e r t a i n extent.  In general i t would seem to lend  support to a more p h y s i o l o g i c a l approach to the e f f e c t s of n e u r a l damage on emotionality.  - 37 -  This approach as employed i n Experiment I seems to have produced data c l e a r l y c o n t r a d i c t i n g the concept of s e p t a l hyperemotionality.  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Neuro-Psychopharmacology. 12, 1967, 427-430. Wood, CD. Behavioural changes f o l l o w i n g d i s c r e t e l e s i o n s of temporal lobe s t r u c t u r e s . Neurology. 8, 1958", 215-220.  -4-3-  APPENDIX  A  Histological reconstructions subjects  o f s e p t a l a r e a damage i n l e s i o n e d  i n experiments I.and I I . Four r e p r e s e n t a t i v e  p e r b r a i n a r e shown 6.2, 7 . 0 , 7 . 8 , and 8.6 mm.  sections  a n t e r i o r to  bregma r e s p e c t i v e l y .  x  J  A  8.6  Septal 1  (  •  '  A 7.8  i  1  i  .  4  I  1 0  1 1  1 2  1 3  I  1 4  5  I  _] 6  I 7  8  f  "  •  ,  '  •  '  I  • I  +  7  r  A  +G  8  6  Expt. 1 Septal 5  <  •  A  8 .6  Expt. 1 Septal 8  A  8.6  Expt. 1 Septal 9  +2  A 6.2  Expt. 2 Septal 1 >.  H r  'I  A 7.0  Expt. 2 Septal 2  A 8 6  Septal 2  +6  A 6.2 +5  Expt. 2 Septal 3  + 4 h  1  «  (  -  •  -  -1  I  + 7  r  A 7.6  Expt. 2 Septal 3  •K7  A  8.6  Expt. 2 Septal 3  + 7  A  8.6  Expt. 2 Septal 5  +  7  A  r  Expt. 2 Septal 6  7.8  6  i  - 5  A 7. 8  8  

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