The Role of the Septo-Hypothalamic Tract and the Nucleus Accumbens in the Septal Rage Phenomenon
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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