This dissertation has been microfilmed exactly as received 70-6822
LAUBER, John K., 1942- THE ROLE OF THE SEPTO-HYPOTHALAMIC TRACT AND THE NUCLEUS ACCUMBENS IN THE SEPTAL RAGE PHENOMENON.
The Ohio State University, Ph.D., 1969 Psychology, experimental
University Microfilms, Inc., Ann Arbor, Michigan THE ROLE OF THE SEPTO-HYPOTHALAMIC TRACT AND' THE NUCLEUS ACCUMBENS IN THE SEPTAL RAGE PHENOMENON
DISSERTATION
Presented in Partial Fulfillment of the Requirements for the Degree. Doctor of Philosophy in the Graduate School of The Ohio State University
By
John Kc Lauberp BeSc,f M»A*
* it * *
The Ohio State University 1969
Approved by
Adviser 7\ Department of Psycholqg] ACKNOWLEDGMENTS
The author wishes to express his appreciation to Professor Donald R„ Mayer9 adviser» for his support and guidance throughout the course of the present study0 I wish to thank Professor Delos Do Wickens and Professor Neal Johnson8 members of my reading committee9 for, their helpful comments concerning the design and write-up of this dissertation0 Thanks also to D r 0 Patricia Mo Meyer for her helpful suggestionss and her assistance during the histological phase of the studyo And f5.nallyp to my wifes Susan M 0 Lauber# for her assistance throughout the entire course of thi dissertations I express my deepest appreciation,.
This research was supported in part by funds supplied through Grants MH-06211 and MH-02035 from the
National Institute of Mental Healths United States
Public Health Service, and was conducted while the author was a predoctoral fellow supported by Grant
MH-06748, VITA
December 13 , 1942 0 6 6 Born - Archbold, Ohio
1965 o o o o o e a b o BoScop The Ohio state University Columbuss Ohio
1965-1969 o c e e o e United States Public Health Service Predoctoral Fellow* Department of Psychology, The Ohio State University* Columbus * Ohio
1967 0 6 0 o a o o o o MoAop The Ohio State University* Columbus* Ohio
1968-1969 o o o a « a Teaching Assistant, Department of Psychology, The Ohio State University, Columbus, Ohio
FIELDS OF STUDY
Major Fields Comparative and Physiological Studies in Brain Mechanisms and Behavior« Professor Donald R0 Meyer and Dr0 Patricia Meyer TABLE OF CONTENTS
Page ACKNOWLLDGMENTS eeoooooesoooooooo 11
VITA o eeeoeeeeoooosoooeo. ooo ilL i
LIST 01* TABLES oosoeooooooeoeooo V
L I S T 01* FIGURil, S o o o o o o o o o o o o o o o o o V I
Chapter
lo INTRODUCTION oooooooeooeoooo 1
Anatomy Background
I I o METHOD QoeOC9Q0OCOO6OeOOO
Subjects Procedure Surgery Histology
III. R ESU LTS eooeoooooeooooooeo lA
IVo DISCUSSION otooooeoooseooco 20
Vo APPENDIX
A oooooooooooooooooooo 23
Boooooooooooooooooooo 23
VI0 BIBLIOGRAPHY ooooooeoooeooeo 26
Iv LIST OF TABLES Page Summary of Mean Emotionality Ratings - Day I Post“Operative e***********. 17
Number of ”Hotf*,,f Medium" , and ,,Cool,, Ss by Groupsj Day 1 Postoperative Ratings* . * 18
v LIST OF FIGURES
Page 1« Mean Emotionalit}' Ratings by Days . o • •> » » 15
vi INTRODUCTION
Brady and Nauta (1953) were the first to describe
in detail the phenomenon known as the septal syndrome0
These investigators produced large,, bilateral lesions
in the septal nuclei of rats 8 and noted subsequent changes
in affective behavior best described as ,!hyperemotionality!' or lshyperj.rtitability” 0 Since this original paper, many
independent investigators have confirmed the Brady and
Nauta observations (Brady and Nauta„ 1955? Kings 1958?
King, 1959? King and Meyer* 1958? Yutzey* Meyer and Meyer*
1964? Krieclchaus et al* 1964? and others)»
One direction taken in some recent research on the
septal syndrome has been that of trying to differentiate the septal complex into components which are responsible for the septal syndrome0 The present research is directly concerned with this problem of differontiation0
Anatomy of the Septal Comolex
The septal area of the forebrain is a complex collection of nuclei and fiber tracts* and forms a major part of the limbic system of lower mammals (MacLean* 1949)„
In this section we shall discuss the major nuclei and fiber tracts* and their interconnections * which form this part of the brain0
1 2
The lateral and medial septal nuclei are the most
prominent of the septal nuclear massest The largest of
these structuresp the lateral septal nucleus0 receives
efferents primarily from the hippocampus via the pre-
commissural system of the fornixs and projects afferent
fibers to lower brain stem structures (Andy & Stephanp
1961)e The opposite relationship has been demonstrated
for the medial septal nucleus9 which receives its major
inputs from lo'wer brain stem structures8 and projects
fibers to the hippocampus (Andy &. Stephan0 1961), Pro
jections from the medial septal nucleus to the amygdala
have also been described (Knoolcp 1965)0
Additional nuclei which are located inp or in close
proximity to? the septal area include the nucleus of the
diagonal band of Brocap the bed nucleus of the stria ter-
minalisp and the nucleus accumbons® Knook (1965) has
demonstrated that the nucleus accumbens receives inputs
- from the hippocampusf the amygdala (via the stria ter-
minalis)p and from olfactory structuresp and sends fibers
to the medial septal nucleus 9 the globus pallidusf and to
various thalamic nucleio
The major fiber tracts which are associated with
the septal complex are the diagonal band of Brocap the
stria terminalisp and the septo-hypothalamic tract® The
stria tarminalis is the most prominent of these fiber
tractsp and has been shown to arise in the basal amygdaloid nuclei, and to terminate in the septum, the pre*»optic areas and the hypothalamus (Fox, 1940, 1943; Halls 1963;
and Knook, 1965) 0 In additions amygdalo-petal fibers
arising from the pre^optic area have been observed in the
stria terminalis (Knook, 1965) ,
Other major tracts which are adjacent to, but not generally considered a part of, the septal area include the corpus callosum, fornix, anterior commissure, and the medial forebrain bundle0
In the usual "septal" preparation, the lateral and medial septal nuclei are almost always destroyed. Total destruction of the nucleus of the diagonal band of Broca, the diagonal band of Broca, the septo-hypothalamic tract, the corpus callosum, and the fornix is only sometimes observed, while partial damage to these structures often occurs. The stria terminalis and its bed nucleus, the nucleus accumbens, the anterior commissure, and the medial forebrain bundle are rarely damaged to any appreciable extent in these preparations.
Background of the problem of differentiation
Brady and Nauta (1953) suggested that the septal rage phenomenon might not be a function of damage to the septum itself *> they noted a tendency for the rage res ponse to be associated with damage to the columns of the fornix. More recent work has failed to confirm this hypothesis, Harrison and Lyon (1957) tried to correlate the septal syndrome with damage to other nuclear masses
and fiber tracts in the septal complex, and failed to
find any significant relationships Bernstein and Brody
(1961) also reported no correlation between nuclear
damage and the septal rage phenomsnon0
Two recent studies may have begun to narrow the
problem of differentiation of the septal complex® Turner
(1969) utilized a stepwise muitiple regression analysis
to discover which of five main components of the septal
area contributed most to the septal rage phenomenon®
Forty-one rats were given bilateral electrolytic lesions
in the septal area following three days of pre-operative
emotionality rating using a modified King (1958) rating
scaleo Following surgery and a recovery period (18 hours)
the Ss were again rated for emotionality® Histological examination of the brains of these Ss yielded an estimate of the per cent destruction of each of five components of the septal areas (l) Nucleus accumbens and anterior eommis sure? (2) Medial septal nucleus and tract of the diagonal band? (3) Lateral septal nucleus? (4) Medial pre-optic area? and (5) Pre- and post-commissural fornix, including the nucleus triangularis septi and the nucleus septalis fimbrialiso The regression analysis was done using the estimate of per cent destruction of these areas as the independent variable, and the difference between the day post-operative ratings as the dependent variable0
Significant (p £ 0 05) regression coefficients were found to be limited to nucleus accumbens damage cn seven of the
12 subscales (Cataplexy* Resistance to captures Resistance to handlings positions Vocalizations Tap responses Puff-2) / and additional!}’’* to the Total Score (sum of ratings on all sub-scales)o Lesions of the lateral septal nucleus were related to changes in one of the sub-scales (Activity) and finally* two of the sub-scales yielded significant regression coefficients to a combination of lesions
(Nucleus accumbens* lateral septum* medial pre-optic area* and medial septum) 0 In summary* damage to the nucleus accumbens* not to the septal nuclei* was found to be con sistently associated with ’•septal1' rage0
The second of these two studies utilized a similar* but more comprehensive* technique0 Meyer et al.(1968) also used a stepwise multiple regression program to assess the,contributions of 38 different nuclei and fiber tracts in and around the septal area to the septal syndrome0
Two experienced investigators independently assessed the brains of 322 rats which had sustained lesions in the septum* combination lesions of the septum and anterior neo- cortex* septum and posterior neoeortex* septum and anterior cingulate gyrus* septum and the entire cingulate gyrus* for damage to each of the 38 nuclei and included fiber tract or nucleus was used as the criterion for inclusion
of a particular brain in a given group of Ss„ These results were then used as the independent variables in a multiple regression analysisj, with Day 1 post-operative emotionality rating scores (using the Kings 1958 emotion ality rating scales) serving as the dependent variables®
The results of consequence for the present study showed that damage to septo-hypothalamic tract (TSiiT) correlated highly with the appearance of septal rage ( p < 0 Ol ) 0
Another structures the pre-commissural system of the for nix,, also proved to be consistently related to the septal syndrome (p <®Ol) 0 but inspection of a matrix of correla tions between var5.ous lesion sites revealed that damage to the pre-commissural fornix was commonly accompanied by damage to TSHT (r=®70)e This observations coupled with those of Harrison & Lyon (1957)P and Turner (1969), sag- gests that it is highly unlikely that the septal syndrome is a function of damage to the pro-commissural fornix®
The present study is an attempt to refine and extend both the Turner and the Meyer et al, experiments® It should be noted that Meyer et al® did not include the nuc leus accumbens in their histological analysis9 since prelimi* nary examination of the brains used in this study revealed that damage to the nucleus accumbens was relatively rare9 and that when it did occurs damage never exceeded the
50% criterion® Neither of these studies'involved Ss 7 with damage restricted to only TSHT or NA; damage to these structures was always accompanied by damage to a large number of adjacent structures0 We have produced rela** tively discrete, localized lesions in both TSHT and NAS and in appropriate control groups„ and have assessed the effects of these lesions upon subject emotionality0 METHOD
Subjects
One-hundred seventy two adults males albino rats
(Sprague-Dawley) weighing between 300 and 450 grams served as ,Ss for this’experiment Ss were housed in individual cages in the laboratory colony room for the duration of the experiment and were fed and watered on an ad libitum basiso
WProcedure in I TT iii 'rilimm— i1 n 111
Ss were rated for emotionality for six days using the emotionality rating scale developed by King (1958)C
An emotionality rating for a given subject consists of the -sum of the ratings given on each of the six sub- scales,, done in the following orders (l) Object presen tations (2) Response to tap on back? (3) Resistance to capture; (4) Resistance to handling; (5) Vocalization to capture and handlings and (6 ) Urination and defecation in reaction to handling., The first five of these scales are scored on a 0-5 basis 8 and the'lasts Urination and defecation,, on a 0-2 basis (see Appendix A for a copy of the scale)„ Simultaneous ratings were done by two inde pendent and experienced raters; the reported emotionality scores represent the mean of the two ratings obtained0 9
On the day following the sixth day of pre-operative ratingc Ss were assigned to one of the four experimental or control groups? (1) TSHT (Septo-hypothalamic tract)? this was our primary experimental groups and its inclusion was based directly upon the results of the Meyer et al 0 computer study? (2) MFB (Medial forebrain bundle)? this group can be viewed as a control group for the TSHT group0
The medial forebrain bundle is also a fiber tractt and, like TSHT* carries both septo-fugal and septo-petal fibers to and from the hypothalamus3 (3) NA (Nucleus accumbens)? this is another nucleus of the septal complex, and was included in the present study as a replication and refine ment of the Turner (1969) study? and (4) LS (Lateral septal nucleus)? this group served as a specific control for the nucleus accumbens lesions9 and in a more general sense for the TSHT lesions as well, since the lateral septum is one of the nuclei (along with the medial septal nucleus) which is damaged in the usual septal preparation0
Bilateral lesions were made in all Ss» Following surgery, Ss were returned to their home cages for two days of post-operative recovery, at the end of which six days of post-operative emotionality rating commenced0
All post-operative ratings were done '‘blind", i0 e 0, neither rater knew to which group a given subject belonged,,
This was insured by having an independent E, who was not one of the two raters, re-number all cages following 10 surgery and then replace these cages in the racks in any order he chose0 A key to the re-numbering was made and kept by this independent E until the end of the post operative ratingo Following the last day of emotionality rating, Ss were sacrificed and perfused intraventricularly with saline and formalin (l07o)o The brains were then removed and' embedded in cello id in for histological analysis.*
Surgery
All surgery v?as conducted as follows? Ss, deprived of food for the 24 hours immediately preceding surgeryt were anesthetized vising a standard, dose of 0 o25 sodium pentobarbital (Nembutal) administered interperitoneally
(XP)o Atropine (Odcc, IP) was used as a pre-anesthetic agento The rats5 heads vie re then shaved and scrubbed using Hexagerm and tincture of ZephiranB E^s were next placed in the stereotaxic instrument, the scalp incision and reflection were made* and a small bone defect was made vising a motorized trephine and rongeurs0
The electrode, 0o020" stainless steel wire coated with insulating Epoxylite except at the 005 mm„ tip, was lowered to the appropriate coordinates (Da Groot), connected to a Grass LM~3 Radio Frequency (RF) lesion maker, and used to deliver 20 seconds of RF current with an anal electrode completing the circuit,. The electrode v?as then withdrawn, swabbed clean with a saline-soaked swab, and lowered into the brain again to complete the 11
bilateral lesion0 The incision was then sutured* 0sl0 cc -
of Penicillin G (Bicillin) was administered intramuscu
larly 9 and the subject was returned to his home cage0
A constant intensity setting (of 1150ts on the arbi
trary lesion-maker intensity control scale) was used for
all lesionso Current was monitored using a VTVM connected
to the lfcurrent monitor11 terminals provided on the lesion-
maker* Current thus measured ranged from 107 ma 0 to 2*8
ma« 5 with a mean current of approximately 2*2 ma 0 This
current* however* is not the electrode tip currenty but
rather, represents the sum of electrode tip current and
the RF current flowing through the capacitance provided by
the interconnecting wires* the electrode holders the
electrode itself* etc* (To measure electrode tip current
accuratelyp a caloric method would have to be used* e 0 g 0
a thermocouple - a far too cumbersome method for the
present purposes,,) All attempts were made to keep these
capacitive losses at a constant minimum5 and the current
monitor was used primarily as an all-or-none indication
that the lesion-making system was functioning properly«
Current parameters (20 seconds at H50!*) were chosen on the basis of a short pilot study* using both behavioral
(iee0y at least some "hot" subjects) and histological
criteria (iee*, damage restricted primarily to the
structure in question),, 12
Histology
After the brains were embedded in celloidin, they were sectioned at 30 rnicrao Two sets of slides were taken from each brain with every fourth and fifth section being retained throughout the lesion site,, One set of sections was stained with cresyl violet Nissl stain (a cell body stain) 5 the second set received a Kluver“Barrera stain
(a combined cell-fiber stain)«
The slides were then examined microscopically0
Drawings of the lesion, including degeneration surroun- ding the lesion site, were placed directly onto the appro priate labeled diagrams Xeroxed from a stereotaxic atlas
(Konig and Klippel, 1963)e Thus, for each brain, a packet of serial diagrams showing the extent of the lesion, and the structures damaged, was obtained,, All further histol ogy was done from these diagrams,,
Ss whose brain diagrams showed no observable lesion were discarded from further consideration, as were those with completely asymmetrical lesions„ For each remaining
S a checklist indicating all structures which had sus tained 50% or greater damage within the anterior-posterior extent of the lesion was constructed,, These data were used to redefine lesion groups as reported in the next section of this paper,. Additionally, a measure of the relative volume of each lesion was constructed by multi plying the anterior-posterior extent of the lesion, as determined from the diagramsp by the cross-sectional area
of the lesion at a point mid-way between its most anterior
and most posterior extents The cross-sectional area was determined by placing a transparent grid marked into one
cm 0 x one ciru squares over the appropriate lesion diagram
and then estimating the area (in square cm,) of the lesion
as shown on the diagram* It must be stressed that this
procedure provided a relative,, not absolute,, estimate of
the volume of each lesion,, RESULTS
The results of both pre° and post-operative emotion ality rating appear below in Figure 1.
Pre-operatively, there were no differences across groups in the emotionality ratings (e.g., Day 6 means were
4.2, 4 0 3, 4.3, and 4.3 for TSHT* -MF3, HA* and LS Ss respectively)fi hence the data shown in the left half of
Figure 1 are the pooled results of all Ss (N=l72)0
A small* gradual decline in ratings over the six day pre operative period was observed. This finding is consistent with the results of previous studies which have involved emotionality ratings of normal animals (e 0 g., Yutzey et al.
1964).
A total of 23 Ss were eliminated from further consid- eration on the basis of either no observable lesion (N=9), or gross asymmetry of the lesion (N=sl4) 0
The right half of Figure 1 presents mean post operative emotionality ratings for Ss grouped according to actual damage sustained as determined by the histo logical analysis. Group TSHT, for example, includes all those Ss which sustained 50% or greater damage to the septo-hypothalamic tract. It should be noted that this re-defined TSHT group can and does ind.ude Ss with other ST N=L5 Hot
Medium N=ll TSHT N=34 NA N=33
Cool
MFB N= 1 0
4,0j All Ss N=l72
Pre-operative days Post-operative days Figure 1. Mean Emotionality Ratings by Days Cn 16
lesions as well, although the amount of overlap is quite
small® Specifically, there are six Ss common to both
TSHT and NA groups® With the exception of the ST group discussed below, all other groups were “pure", e 0 go, MFB
contains no TSHT Ss 9 etc 0
The group designated ST was included in the present
analysis when an unexpectedly large apparent correlation
between emotionality and damage to the stria terminalis
(ST) and/or its bed nucleus appeared in the histological check listso There is a relatively large amount of over
lap of five Ss which are common to ST and TSHT, and of three Ss common to ST and NA« There are no higher-order overlaps however, i®e0, those Ss sustaining both NA and
ST damage do not have TSHT damage as well, nor do TSHT-ST
Ss have NA involvement®
Table 1 presents a summary of Day 1 postoperative emotionality rating data® Comparison of the mean emotion- ality rating of each group with the mean rating of all other S_s combined indicated that MFB animals scored significantly lower than all other S_s0 On this basis, and on the basis of lesion placement (lateral and poster ior for MFB lesions, medial and anterior for TSHT* LS* and
NA lesions), MFB Ss were removed from further consideration in Table 1 0 Thus, MFB animals do not appear in any of the pooled-groups columns of Table 1® 17
Table 1„ Summary of Mean Emotionality Ratings » Day 1 Post-Operative
tf~-~jseaag»rg:*.ari«Mi iijm L i'UJCT w s r .ii'-LUU*a«nP»rasg»rTi»»'jj>m» g !tg ^ «-A.’
MFB 1 0 6 00 All Others 139 8*9 2.41/47 < * 025
TSHT , 34 9*6 o A All Others 76 9 o0 <1/108 o
NA 33 9.1 All Others 77 9c 3 <1/108 > e 05
LS 1 1 lOo 8 All Others • « 99 9*1 lo30/108 > *05 ST 15 14 * 4 00 Ln All Others 95 Q 6.21/108- < o 005 *A11 probabilities based on one-tailed t distribution T aw iB.-»-saagijaa3aTs-*ng«^;uiK»ofrgngisa*>»
Comparisons between the mean emotionality of NAp
TSHT, and LS Ss indicated no significant differences between any of these groups and all other _Ss combined, nor between any two specific groups (all p> o05i see
Table 3, Appendix B for a summary of all group compari sons) „
Animals with lesions involving ST scored signifi cantly higher in emotionality ratings than did all other
Ss combined, and were significantly ••hotter” than any other specific group ( p < o005)o
Product“moment correlation coefficients relating i lesion size (based on the relative volume of damage estimate outlined previously) were all small ( < * o25), and did not approach statistical significance (p> »1 0 )„ There is no evidence of a mass-action effect in these data*
Table 2 shows the results of breaking down the rating scales into three intervals which can be consid ered roughly as "cool"s "medium" B and f!hot" subjects., The intervals showne 0~7o9 (cool)p 8o0-llo9 (medium) 9 and 12*0-
27*0 (hot) 9 were arrived at independently and a priori9 by
*1 tliree experienced raters 9 and represents their best esti mate as to how to divide the King (1958) scale into the three categories shown* These intervals are also indi cated on Figure 1 by the horizontal lines drawn from the appropriate points on the ordinate*
Table 2* Number of "Hot", "Medium" 9 and 11 Cool1' Ss by Groups! Day 1 Post-Operative Ratings
Emotionality Ratings Group N 0-7*9 8 . 0 - 1 1 . 9 12o0-27 o 0
MFB 1 0 7 3 0
NA 33 1 6 8 9
LS 1 1 3 2 6
TSHT (less ST) 29 1 2 9 8
ST 15 1 1 13
t-|-rr TTTiTrrii ir-w n» -r n r riiwwi
As indicated in Table 2S and in Figure 1, MFB lesioned
3 s almost always fall into the "cool" category» and none lDr„ P* Me Meyer 9 Susan M# Lauber 9 6c Dr* David A* Yutzey of these Ss were ”hot”„ ST animalsp however? are almost universally ”hot”, while TSHT? W A S and LS Ss appear to be more evenly distributed among the three categories..
It should be pointed out that the row in Table 2 indicated by "TSHT less ST” represents the result of removing the five overlapping ST Ss from the TSHT group,, an operation which lowers the mean of the remaining Ss from 9e6 to 8e9, but leaves the relative standing of the group unchanged with respect to LS and NA Ss (see Table 3? Appendix B)® * DISCUSSION
The present results do not support the Meyer et al„
(1968) hypothesis that the septo-hypothalamic tract is / ■ the crucial locus for the septal rage syndrome, nor the
Turner (1969) hypothesis that the nucleus accumbens is
the critical structure,, Although "hot11 subjects are found
in groups of rats with lesions In either of these struc tures, they do not appear with any great degree of consis- tency. In fact, statistically, neither of these groups
is •'hot111 the mean Day 1 post-operative emotionality ratings for both groups fall within the •'medium*1 category0
It is conceivable that the relatively high correla tions between emotionality and damage to TSHT and NA observed in the Meyer et al. and Turner experiments can be attributed to the close proximity of each of these struc tures to a third structure, the stria terminaliso The present data strongly suggest that damage to the various components of ST is of major importance with respect to the appearance of septal rage. Turner, in a recent sequel to his earlier experiment, lends support to this notion,. Tentative results of this experiment, which, like the present study, utilises small lesions within the septal area, indicate that septal hyporernotionality
20 21
is related to the stria terminalis (Turner, 1969) 0
Anatomical and physiological data involving the
stria terminalis are consistent with the hypothesis that ST damage is related to the appearance of septal rage0 Fox (1940, 1943), Hall (1963), and Ban & Omukai
(1959) have shown that the stria terminalis is composed of fibers which are predominantly efferent with respect to the amygdala, and which distribute from that struc ture to the septum, hypothalamus, and the pre-optic area#
Lesions of the anygdala are known to produce both ucooln and "hot** animals (Cfo Kleiner et: al, 1967), and electri cal stimulation of the same structure has been shown to modulate hypothalamic functions# Egger & Flynn (1967) stimulated in various parts of the amygdala atid observed the effects of such stimulation upon attack behavior elicited by concurrent stimulation of the hypothalamus#
When amygdaloid stimulation was delivered to the basal and medial portions, a supression of ongoing attack was observed, whereas a lesion in this same area resulted in a facilitatory effect# Knook (1965) has observed degen eration in generally basal and medial portions of the amygdala following lesions of the stria terminalis, which implies that the principal origin of these fibers is within the same general area of the amygdala that Eggers and Flynn have shown is capable of modulating hypothalamic
'•rage1’ responses# In view of the present results* and the anatomical and physiological data discussed above, an hypothesis relating septal hyperemotionality and the stria terminalis is appealing* but will be completely resolved only by further research* APPENDIX A
SIX RATING SCALES FOR THE STUDY OF EMOTIONALITY lo REACTION TO OBJECT PRESENTATION s Pencil is pre sented close to the animal8s snout* 0- Rat ignores pencil* 1- Rat alert and attentive, some body tenseness* 2- Legs and body tense and immobile,, vibrissae point forward* 3- Scurries away or makes 'occasional mild biting attack on pencilo 4- Intermediate* 5~ Very aggressive attack,, disorganized panic, or violent flight*
II. RESPONSE TO TAP ON BACK 0- No reaction 1- Twitching or restlessness* 2- Twitching or scurring away* 3“ Jumps or hops up a bit, but then settles down* 4 “ Leaps in air and turns about in fright* Big hop and movement after* 5“ Leaps violently, dashes off in panic, frantic rebounding from side to side of cage*
III* ’ RESISTANCE TO CAPTURE! Glove is extended forward to animal slowly and rat is grasped firmly but not roughly* 0- Remains calm, does not move when approached, does not struggle when grasped* 1- Remains calm when approached', but scurries away and tugs a bit when grasped* 2- Avoids on approach, struggles vigorously when grasped* 3- Retreats when approached, struggles vigorously when grasped* 4- Strong attempt to escape when approached, struggles strongly and is disorganized, - some biting* 5- Leaps violently when grasped, bites frantic ally, exceedingly difficult to catch*
23 RESISTANCE TO HANDLING 0- Relaxes in hand* does not attempt to escape# 1~ Restless with some feeble squirmingo 2- Sporadic attempts to pull out of hand 0 3- struggles pretty continuously and is quite vigorous in attempt to escape0 4~ Bites also# 5- Frantic biting* powerful tugging and disorganized twisting,,
VOCALIZATION TO CAPTURE AND HANDLING 0“* None, 1“ Few squeaks# 2- Frequent squeaking# 3= Frequent squealing“squawking# 4» Squealing continuously. 5*» Frantic and loud screeching continued#
URINATION AND DEFECATION IN REACTION TO HANDLING 0“ None# 1“ Urine 1 #2 “ 1 stool 1#5° Urine and 1 stool 1 #S~ 2 stools 2c0~ Urine and 2 stools APPENDIX B
Summary Table 3. t tests
Comparison t df p*
ST vs All Others 6 o 21 108
*All t tests* one-tailed probabilities
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