The Effects of Diagonal Band of Broca Lesions Upon Pain-Elicited Fighting and Water Intake

Western Michigan University ScholarWorks at WMU

Master's Theses Graduate College

12-1971

The Effects of Diagonal Band of Broca Lesions upon Pain-Elicited Fighting and Water Intake

David A. Nolley

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Recommended Citation Nolley, David A., "The Effects of Diagonal Band of Broca Lesions upon Pain-Elicited Fighting and Water Intake" (1971). Master's Theses. 2915. https://scholarworks.wmich.edu/masters_theses/2915

This Masters Thesis-Open Access is brought to you for free and open access by the Graduate College at ScholarWorks at WMU. It has been accepted for inclusion in Master's Theses by an authorized administrator of ScholarWorks at WMU. For more information, please contact [email protected]. THE EFFECTS OF DIAGONAL BAND OF BROCA LESIONS UPON PAIN-ELICITED FIGHTING AND WATER INTAKE

David A. Nolley

A Thesis Submitted to the Faculty of The Graduate College in partial fulfillment of the Degree of Master of Arts

Western Michigan University Kalamazoo, Michigan December 1971

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGEMENTS

The author would like to express sincere appreciation to

Dr. Frederick P. Gault, chairman of the thesis committee, for his

instruction and advice during the entire course of this thesis.

Thanks are also to be given to Dr. Bradley Huitema and Dr. Ronald

Hutchinson, the other members of the thesis committee, for their

suggestions and constructive criticism . The author is also

indebted to Miss Leta Hunt for help with the reliability measures

and to Mrs. Judy Buelke for assistance with the final preparation

of the manuscript.

finally, I would like to thank my wife, Barbara, for her

assistance, constructive criticism and patience during every portion

of the thesis.

David A. N olley

i i

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. MASTERS THESIS M-3214

NOLLEY, David Allen THE EFFECTS OF DIAGONAL BAND OF BROCA LESIONS UPON PAIN-ELICITED FIGHTING AND WATER INTAKE.

Western Michigan University, M.A., 1971 Psychology, exoerimental

University Microfilms, A XEROX Company , Ann Arbor, Michigan

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS

CHAPTER PAGE

I ACKNOWLEDGEMENTS...... i i

I I TABLE OF CONTENTS ...... i i i

I I I LIST OF TABLES AND FIGU RES...... iv

IV INTRODUCTION ...... 1

V METHODS AND PROCEDURES ...... 3

S u b jects ...... 3

Surgical Procedures ...... 3

Apparatus and M aterials ...... 5

Dependent Measures ...... 7

VI RESULTS ...... 8

Affective Behavior ...... 8

Water Intake ...... 11

Histological Verification ...... 12

VII DISCUSSION...... W-

V III FOOTNOTES ...... 21

IX BIBLIOGRAPHY...... 22

FIGURE CAPTION...... 27

i i i

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF FIGURES AND TABLES

PAGE

TABLE I ...... 18

FIGURE I ...... 19

TABLE I I ...... 20

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. INTRODUCTION

There is a marked difference in affective behavior resulting from

lesions of the septal area and the amygdala. Septal lesions have

generally resulted in hyperirritability and hypersensitivity to

preoperatively neutral stimuli (Brady and Nauta, 1953; 1955; King, 1958)

Conversely, the weight of evidence has suggested that lesions of the

amygdala produce a relatively docile, hyposensitive animal (Ahmad and

Harvey, 1968; Schreiner and Kling, 1953; Rosvold, Mirsky and Pribram,

1954; Weiskrants, 1956; Shealy and Peele, 1957; King, 1958).

With respect to water intake, Grossman and Grossman (1963)

reported that hyperdipsia resulted from posterior-ventral amygdaloid

lesions. The effects, however, were not permanent unless the lesions

were quite substantial. In the same study, electrical stimulation of

the anterior amygdala increased water consumption, while stimulation of

the posterior amygdala depressed water intake. Lesions of the medial

amygdaloid nuclei resulted in profound adipsia (Collier and Gault, 1969)

Grossman (1964) reported increased water drinking with chemical

stimulation of the septal area. However, several authors have reported

hyperdipsia after septal lesions (Harvey and Hunt, 1965; La Vaque, 1966;

Kasper-Pandi, Schoel and Zysman, 1969) and suction ablations (Lubar,

Boyce and Schaefer, 1968).

After Broca's ( 1878) description of the diagonal band as a part of

the limbic lobe, a number of authors have referred to it as a fiber

pathway between the amygdala and the septal area. Johnston (1915* 1923)

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Loo (1930. Young (1936), Humphry (1936) and Fox (19^+0) traced, from the

medial septal nucleus, medial diagonal band fibers to the anterior

nucleus of the amygdala and a lateral component of the band to the

globus pallidus. Fox (19^+0) notes that as the diagonal band of Broca

(EBB) passes through the medial forebrain bundle, some of its fibers

separate to compose the nucleus pre-opticus magnocellularis. The band

then continues, ventral to the medial forebrain bundle, on to the

amygdala.

The pronounced behavioral differences resulting from lesions of the

septal area and the amygdala, together with the hypothesis that the DBB

provides interconnections between the two areas, points to the proposi

tion that the diagonal band may convey some influence of one of these

structures over the other. This study presents data concerned with

correlated changes in affective and water intake behavior resulting from

lesions of the DBB and septal area. Pain-elicited fighting (Ulrich and

Azrin, 1962) was used to measure changes in affective behavior, while

daily water intake measures were made. Each pair of animals was used as

its own control, for the data reported.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. METHODS AND PROCEDURES

S ubjects

Thirty male, experimentally naive Sprague-Dawley rats, weighing

between 180 and 310 grams at the initiation of the study, served as

subjects. The animals were randomly assigned to pairs, concurrent with

the requirements of the pain-elicited fighting paradigm. Data are

reported for only fourteen of the fifteen pairs because severe post

operative debility was observed in one member of one of the pairs.

Changes in water intake are reported for twenty-three of the subjects.

The animals were housed individually and were allowed free access to

ad lib food and measured quantities of water.

Surgical Procedures

All surgeries were conducted under clean but not asceptic conditions

using sodium pentobarbital (40 mg/kg intraperitoneal) anesthetic.

Atropine sulfate was used to reduce respiratory secretions. Subjects

were placed in a Kopfl stereotaxic instrument consistent with the

procedure described by Konig and Klippel (1963). Burr holes were made in

the skull at the appropriate coordinates for the specific operation after

a midline incision was made and the periostium retracted. In all cases,

bilateral lesions were produced electrolytically using stereotaxically

placed electrodes (Clay-Adams2 #00 insect pins, insulated with Formvar3,

except for 0.5 am at the tip), passing anodal DC through the uninsulated

tip of the electrode. The lesion circuit was completed by a reference

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electrode.clipped to the ear bar.

For septal lesions, an angular approach was utilized, using the

coordinates: anterior 7«1 mm, lateral 0.6 mm and 0.8 mm above zero

(Konig and Klippel, 1963). Septal lesions were made by passing a

current of 2.0 ma for thirty seconds. For DBB lesions an angular

approach was employed to the coordinates: anterior 8.0 mm, lateral

0.2 mm and 2.0 mm below zero. A current of only 1.0 ma for 25 seconds

was used because of the smaller size of the diagonal band relative to

the septal area. For sham operates, electrodes were placed in the

appropriate structure without passing current and then withdrawn.

Since the Konig and Klippel atlas was constructed upon rats

weighing 150 grams, a correction for growth of the brain was made for

the heavier rats used in this study, for coordinates which exceeded

1.0 mm from atlas zero. For rats weighing 200-245 grams at the time of

surgery, 10$ was added to the appropriate coordinates. For rats

weighing from 250 to 305 grams, 15$ was added. Animals weighing 310

to 385 grams required a 20$ correction. Animals weighing 390 to 445

grams required a 25$ correction. For those animals weighing over 450

grams, 30$ was added to the appropriate coordinates.

In an cases, both animals in a pair were operated upon after an

appropriate number of daily sessions was allowed for a baseline of

behavior. Eighteen to thirty hours were allowed for recovery before

the first post-operative measurements were made.

From five to twenty-three daily sessions were run previous to the

first surgery, for normal pre-operative baselines. In the case of

sequential procedures, seven to fourteen sessions were allowed after

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diagonal band lesions and six to eight sessions after DBB sham lesions.

Only the first six to ten sessions after septal lesions were used in

the manipulation of the data, because of the transitory quality of the

"septal syndrome" (Brady and Nauta, 1955; Yutzey, Meyer and Meyer,

1964), However, two of the pairs which received septal lesions as

their second surgery were run up to fourteen sessions after their

surgery. One of these pairs had previously received diagonal band

le s io n s , th e o th e r had received sham DBB le s io n s .

With the exception of four pairs, all subject pairs were assigned

their first surgery by a random method. The assignment of second

surgeries, in the case of sequential procedures, depended upon the

type of surgery given first. All pairs were generally run every day,

approximately every eighteen to thirty-six hours.

At the completion of the experiment, the animals were sacrificed

with an overdose of sodium pentobarbital and perfused intracardially

with saline and 10$ formol-saline. The brains were removed from the

head, sectioned at fifty micra intervals and the lesions verified

histologically using the method of Guzman-flores, Alcaraz and

Femande z-Guardiola (1 963) •

Apparatus and M aterials

Pain-elicited fighting sessions were run in a chamber with an

effective fighting area of 9 1/4x9 1/2x12 inches. The chamber

was positioned approximately three inches from a one-way mirror through

which the experimenter viewed the subjects. Three walls of the chamber

were constructed of 1/2 inch plywood, painted flat black on the inside

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to maximize contrast for photography. The fourth wall was constructed

o f 3/8 inch plexiglas. The floor of the chamber was constructed c*

eighteen parallel 1/8 inch stainless steel rods, approximately 1/2 inch

apart at the centers. Chamber illumination was provided by a fifteen

watt bulb, protected by a wire cage set high upon one of the plywood

walls of the chamber. The hinged plexiglas top of the chamber had ten

1/2 inch holes drilled through it to provide ventilation. Scrambled

foot shock was delivered by a Grason-Stadler^ E60?0B shock generator/

scrambler. Amperage delivered by the shock generator, though

consistently the same during the usefulness of each pair, differed

among pairs to demonstrate that the pain-elicited behavior was not

specific to any particular level of current. Three pairs were run at

0.3 ma. Three pairs were run at 0.8 ma. Eight pairs were run at 1.3

ma.

During each session, 0.5 second shocks were delivered with a 15»0

second intertrial interval for a total of 120 trials, programmed by a

Lehigh Valley5 lh .36 d ig i ta l Countdown Timer, two G rason-S tadler E1100H

Running Time Meters and associated electromechanical circuitry. The

experimenter in the observation room recorded fighting responses and

trials during which fighting took place using hand operated micro-

switches. Fighting responses and shock presentation were recorded on a

Gerbrands^ cumulative recorder.

A duplicate recording system was used for sessions in which

reliability checks were made, with the cumulative recorders measuring

fight responses placed in another room such that there was no auditory

feedback to contaminate the measures of reliability. The two observers

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sat side by side in front of the one-way glass to approximate an

identical view of the animals in the chamber.

Dependent Measures

There were two dependent measures used to record fighting behavior.

One was a simple binary measure: the presence or absence of fighting

responses during each trial of each session, which was then converted

to a percent measure of trials per session during which fighting

occurred. The other was subjective measure of cumulative fighting

responses during each session observed by the experimenter. Sighting

responses were similar to those employed by Bryant (1969). A response

was defined as contact made by the paws of one subject on the other

subject inclusive of an area defined by a line between and just behind

the ears and running diagonally and posteriorly to a point where the

abused ra t's elbow would rest when down and drawn in toward its body.

Contacts were not counted if: a) the defined response area was

contacted with the hind feet of the other subject; b) if the subject

making the attack had one of its front paws on the grid, even though

proper contact was being made with the other paw; or c) if the abused

su b je c t had n o t made a t le a s t one paw swipe a t h is a tta c k e r during

that trial.

Water intake was measured in m illiliters consumed daily and was

measured approximately every twenty-four hours.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. RESULTS

Affective Behavior

Diagonal band lesions consistently depressed fighting rate, as

reflected by the percent of trials measure and the total cumulative

number of fight responses, in all pairs. The diminution in aggressive

behavior was abrupt, occurring during the first session after surgery.

This occurred whether the lesion had been proceeded by a normal

baseline or one heightened by septal lesions. The depression was

maintained throughout the seven to twelve post-DBB sessions used to

establish a baseline prior to septal surgery.

In the pairs given diagonal band surgery after a septal baseline,

the startle response, characteristic of the “septal syndrome* (Brady

and Nauta, 1953)»did not generally diminish until four to five d a i ly

sessions after the band of Broca surgery. Characteristic, however, of

the pairs receiving DBB surgery after a normal baseline, these pairs

consistently attempted to avoid facing each other by either facing a

corner or arranging themselves such that they were facing opposite

walls of the experimental chamber. This behavior was evident from the

first session post-surgery and was observed to occur even after a

strong startle response was elicited by the footshock during the early

portion of the DBB baseline.

Two pairs were maintained on their DBB baseline without the septal

surgery to gauge the long term effects of band of Broca surgery upon

affective behavior. The depression in these pairs was maintained at

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least twenty-one daily post-operative sessions. Between twenty-one

and twenty-five sessions after the diagonal band surgery, both pairs

again fought several sessions at their pre-operative rate. One pair,

then, again approximated its diagonal band baseline while the other

pair remained near its pre-operative rate of fighting.

As shown in Table 1, th e s u b je c t-p a irs given DBB sham le s io n s

after an unoperated baseline showed no significant difference in either

of the dependent measures. Their fighting was unremarkable when

compared with their pre-operative baseline. The dominant-submissive

relationships established during the pre-operative baseline in the

experimental chamber did not change post-operatively, as did a number

of those relationships established in pairs which received septal or

band of Broca lesions.

All pairs given the septal surgery fought on a higher percent of

trials when compared to their normal pre-operative baseline (ns). With

th e exception o f p a ir E, mean cum ulative number o f fig h tin g responses

were also higher (ns) after 3eptal lesions. Normal pre-operative

fighting in these subject-pairs, however, was so high that there may

have been a diminished opportunity for the septal surgery to illustrate

a change in fighting sufficient to reach significance. Since paw

contacts of a specified topography were used as the criterion for

fighting behavior, the enhanced startle exhibited by all septally

lesioned animals for the first few sessions after surgery often

prohibited proper positioning for paw contact in the defined response

area .

As a case in point, septal lesions given after a sham DBB lesion

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 10

baseline resulted in significantly higher percent of trials in which

fighting occurred and cumulative fighting responses when compared with

the sham diagonal band baseline. None of the sham pairs had fought

with either as high a percent of trials or cumulative responses, pre-

operatively, as the pairs given septal lesions after a normal baseline.

In all other respects, the behavior of the animals given septal lesions

after band of Broca sham lesions mimiced the affective behavior of the

animals with septal lesions given after a normal baseline. The

characteristics of pain-elicited fighting prior to and subsequent to

septal lesions have been adequately described by Bryant (19&9)*

Septal lesions given after diagonal band lesions raised both

affective dependent measures to an extent approximating that observed

in subject-pairs lesioned after a pre-operative or sham DBB baseline.

However, as depected in Figure 1, this heightened fighting was quite

transitory, lasting not more than two sessions in two of the pairs and

not more than five sessions in the other two pairs. Both affective

measures then approximated and remained at the DBB baseline until the

animals were sacrificed. Unfortunately, there were insufficient data

to statistically compare this group with an adequate control group.

Measures of the reliability of th^ observation technique were

gathered at least once during each baseline period for each pair of

animals. The Person Product-Moment correlation between the percent of

trials fighting occurred tallied by the two observers was *9?8. The

correlation between the cumulative number of responses observed was .989*

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Water Intake

As shown in Table 2, diagonal band lesions given as a first

surgery, given after a septal baseline, and compared with DBB sham

surgery increased water intake significantly. Only one of fifteen

subjects failed to increase daily water intake subsequent to the band

of Broca surgery. Eight of these subjects increased their intake on

the day following their surgery. Two of these eight subjects had

received their DBB surgery after a pre-operative baseline. In these

animals, water intake began to fall after four days, eventually

approximating the pre-operative intake. The water intake of the other

six subjects, those that had received their DBB lesions following a

septal baseline, did not deviate from the elevated level until they

were sacrificed and perfused.

In six of these fifteen animals, an increase in water intake did

not begin until two to four sessions after the band of Broca surgery.

The intake then remained at this elevated level in three of these six

subjects while it fell to the pre-operative level in the other three

animal3. There was no systematic correlation between type of pre

operative baseline and consumption of water in these six animals.

In seven of twenty-three subjects, there was an increase in water

intake following septal lesions. Five of these seven animals began

drinking at elevated levels by the first post-operative day; the other

two in creased t h e i r in ta k e two to fo u r days a f te r su rgery. Four o f

these subjects continued to drink at an elevated level until they were

either sacrificed or given their second surgery. There was no

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systematic correlation between type of pre-operative baseline and

consumption of water in these seven subjects. The differences between

the pre-operative and DBB sham baselines, the pre-operative and septal

baselines, the DBB sham followed by septal baselines and the diagonal

band followed by septal baselines did not reach significance.

Histological Verification

Lesion placements are not reported for two subjects whose brains

received extensive destruction, presumably as a result of the perfusion

procedure. Partial reconstruction of lesion damage is available for

sixteen of the remaining twenty-eight animals. In four animals,

cortical infection invaded the radiation of the corpus callosum.

In animals with septal lesions only, there was bilateral destruc

tion of the anterior portions of the medial and lateral septal area in

all animals, with posterior septal destruction in two subjects. In

rare instances, the anterior portion of the fornix, the cortical

portion of the columns of the fornix, the precommissural fornix, the

septo-hypothalamic tract, the nucleus of the stria terminalis, the genu

of the corpus callosum, the corpus callosal tract, the triangular

septal nucleus, the hypothalamic tract and the hippocampal commissure

were damaged.

In animals with diagonal band of Broca lesions only the nucleus

and tract of the diagonal band were destroyed in all subjects. In one

or two animals each, the lesions invaded the medial forebrain bundle,

the medial preoptic nucleus, the anterior commissure and the nucleus

of the stria terminalis.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. In animals that received both septal and diagonal band lesions,

destruction of these structures was as described above. Less frequently

involved structures were the corpus callosum, the septo-hypothalamic

tract and the medial preoptic nucleus. In one or two animals each,

there was involvement of the nucleus of the stria terminalis, the

lateral preoptic nucleus, the medial forebrain bundle, the hippocampus

pars anterior, the supra-chiasiuatic preoptic nucleus, the periventric

ular preoptic nucleus, the genu of the corpus callosum, the nucleus

accumbens, the anterior commissure, the superior portion of the fornix

and the columns of the fornix. In three of the subjects, the posterior

lateral septal area was also destroyed.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. DISCUSSION

There are ample data (Schreiner and ELing, 1953; 1956; Weiskrantz,

1956; Shealy and Peele, 1957; King, 1958; Reynolds, 1 965) "to demonstrate

that amygdaloid lesions produce deficits in affective behavior,

particularly when placed in a "social" environment, although Spiegel,

et. a l. (1940), Bard and Mounteastie (1948) and Bard (1950) have

presented conflicting observations. Most of the ablation studies

dealing with the amygdala, however, have involved the entire amygdaloid

complex, a factor which has contributed to the confusion regarding the

influence of this structure upon affective behavior. Studies which

have attempted to delimit more precisely the nuclear masses involved

and correlate the behavioral changes with selective partial destruction

of the amygdala have contributed a great deal towards resolution of

some of the confusion. Green, Clemente and de Groot (1957), after an

extensive analysis of the amygdala, were able to confine their results

which showed hypoactivity and hypoirritability to lesions of only the

anterior amygdala.

One of the two efferent terminuses of the diagonal band of Broca

is in the anterior amygdala (Johnston, 1915; 1923; Loo, 1931; Young,

1936; Humphry, 1936; Fox, 1940). The p re se n t study has shown th a t

diagonal band lesions clearly mimic the effects of anterior amygdala

lesions with respect to the depression of irritability and pain-elicited

aggression, even when this type of affective behavior has been

heightened hy septal lesions. The abject lethargy reported by Green,

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et. al. (1957) after lesions of the anterior amygdala was not observed

in these subjects given diagonal band lesions.

Because of its rich connections with subcortical structures,

several investigators have suggested that the chief function of the

amygdala is a modulatory one (Bard and Mountcastle, 19**8; Gloor, 1955).

Further evidence for this position is suggested here. There is the

implication that the nucleus of the diagonal band of Broca shares the

function of modifying affective behavior with a number of other

subcortical structures including the septal area (Brady and Nauta, 1953;

1955. King, 1958), the hippocampus (MacLean and Delgado, 1953; Naquet,

195*0» the cingulate gyrus (Smith, 19****; Ward, 19**8; Bard and

Mountcastle, 19**8) and the hypothalamus (Hinsey, 19**0; Egger and Flynn,

196**). It is proposed that this mitigating influence of the diagonal

band upon irritab ility or rage is then modulated by the anterior

portion of the amygdala. That the band of Broca lesions did not result

in the "clay-like rigidity" reported by Green, et. al. (1957) after

anterior amygdaloid lesions, gives this interpretation some weight and

suggests that the depression in affective behavior modulated by the

amygdala is influenced by one or a number of other structures aside

from the band of Broca.

Further support for this proposition is provided by Bryant (1969)

and Rosvold, et. a l. (195**) who indicated that lesions of the amygdala

exerted a profound influence upon aggressive behavior. Bryant was able

to reduce pain-elicited fighting in rats 60$ to 90$ after normal pre

operative baselines and heightened septal baselines. Rosvold, et. al.

reported that their amygdalectomized monkeys never emitted aggressive

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responses towards other monkeys in a social situation after their

surgery. The diagonal band lesions in this study rarely reduced pain-

elicited fighting more than 60$ .

In addition, Bryant gave his amygdalectomized subject-pairs

septal lesions, after stabilization of their amygdaloid baseline, with

the result that no differences in these pairs* depressed rate of

fighting was observed. As shown in this study, septal lesions given

after a diagonal band baseline of aggression can again raise the

frequency and intensity of pain-elicited aggression, if only for a

transient period.

In every case, depression of affective behavior after diagonal

band lesions that was not as severe as that observed in most of the

subject-pairs, was correlated with lesion damage which had invaded the

nucleus of the stria terminalis. According to Fox (19^3). the stria

terminalis in cats contains exclusively efferent fibers which arise

from the cortical-medial amygdala and are distributed primarily to the

septal, preoptic and hypothalamic areas. Reynolds (1965) proposes that

the "septal rage" is a function of incidental damage to other structures.

Though the use of affective measures other than emotionality ratings

and checklists have give additional credence to the persistence of

hyperirritability resulting from septal lesions (Wetzel, Conner and

Levine, 1967; Bryant, 1969), there is the implication that the stria

terminalis, as a afferent input to the septal area, assumes either one

of two functions. There is the possibility that it functions in a

primary position in altering affective behavior mediated by the septal

area, or it may carry an influence upon the septal area from those

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portions of the amygdala which have been shown to influence affective

behavior* In either case, lesions which invaded the stria terminalis

may have had an effect upon the affective measures employed in this

stu d y .

Partial reconstructions were made of the lesions of six of the

seven subjects that showed an increase in water consumption after

septal lesions. In five of these subjects, the septal lesions invaded

the posterior portions of the lateral septal area. In none of the

other septal lesions performed, was there damage to the posterior

septal area. These results are consistent with those of Lubar, et. al.

(1968)* Kasper-Pandi, et. al. ( 1968), however, were unable to show a

pattern of damage within the septal area consistently associated with

hyperdipsia, though they consistently produced hyperdipsia with septal

lesions. Their results are consistent with those reported by Harvey

and Hunt ( 1965) and La Vaque ( 1966). In the twenty-three animals in

this study that received septal lesions, however, only the seven

previously cited became hyperdipsic after septal lesions.

It has been shown that the diagonal band of Broca arises in the

medial nucleus of the septal area (Johnston, 1915; 1923; Loo, 1931;

Toung, 1936; Humphry, 1936; ?ox, 19^) • Only one subject of the

fifteen in this study given DBB lesions failed to show an increase in

water intake. It is proposed that, via interconnections with the

medial septal area, the posterior septal area influences the diagonal

band with respect to drinking behavior, and that the effects of septal

lesions upon water intake are primarily those of the posterior lateral

septal nuclei.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Treatment Comparisons obt. t df

Septal lesions followed $ of trials 15.15 3 <.001 by DBB lesions cum. responses ?.76 3 < .01

Preoperative followed $ of trials 7.41 5 < .001 by DBB lesions cum. responses 4.6l 5 < *01

♦DBB lesions compared £ of trials 10.02 8 < .001 to DBB shams cum. responses 2.57 8 < .03

DBB sham followed i of trials 9.01 3 < .01 by septal lesions cum. responses 4.26 3 < .05

DBB lesions followed Assumptions for statistical treatment of by septal lesions this group could not be satisfied

Preoperative followed j> of trials 2.46 > .05 by septal lesions cum. responses 2.13 > .10

Preoperative followed $ of trials 1.74 3 > .1 0 by DBB shams cum. responses .24 3 > .5°

Table 1. Summary of changes in Affective Behavior. Tests are t-tests for difference between means using correlated samples except *, which is test using independent samples

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. without prohibited reproduction Further owner. copyright of the permission with Reproduced

MEAN PERCENT OF TRIALS FIGHTING OCCURRED 01 / I 11000 0 0 1 -1 I / 1001- 20 O CT H400 0 4 H T C O - 1 0 4 — 300 0 |3 - ^ ' |— 0 3 - \ . \ -1500 \ ./ 1 \ |- 0 5 o— . |oo -|io ..O io|— 6 0 |- \ ' ' \ |- 0 6 01- 1 1700 0 7 -1 1 s X - 1 70 0- {800 0 8 -{ 801- - j 1900 0 9 -1 j |- 0 9 | - \ \ - | 0 . O 2 4 5 6 5 4 3 2 1 ------# I / O 1 / O ■ / — ^-o - ^ O— OTSPA SESSIONS POST-SEPTAL ECN O TRIALS OF PERCENT 0 i ' / i ' i l \ \ / \ \ I u \ UUAIE RESPONSES CUMULATIVE 19 # \ T ------B ~|llOO DBB SEPTAL OBB SEPTAL

\ 600 0 |6 - \ \ \ #

| 0 0 -|2 EN UUAIE UBR F EPNE PR SESSION PER RESPONSES OF NUMBER CUMULATIVE MEAN Treatment Comparisons obt. t df

Septal lesions followed 4.54 8 < .002 by DEB lesions

Preoperative followed 2.74 5 < .05 by DBB lesions

*DBB lesions compared 3.26 12 < .01 to DBB shams

DBB shams followed .40 7 > .5 0 by septal lesions

DBB lesions followed .66 5 > .50 by septal lesions

Precperative followed 1.27 8 > .20 by septal lesions

Preoperative followed .1 4 7 > .50 by DBB shams

Table 2. Summary of Changes in Water Intake. Tests are t-tests for difference between means using correlated samples except *, which is test using independent samples.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. FOOTNOTES

^Kopf Instruments, Tujunga, California 2 Clay-Adams Inc., New lork. New York 3 General Electric, Bridgeport, Connecticut 4 Grason-Stadler, West Concord, Massachusetts

^Lehigh Valley Electronics, Fogelsville, Pennsylvania

^Harvard A pparatus Company, Dover, M assachusetts

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. FIGURE 1. Comparison of Means of Fighting

Behavior of Four Septal Pairs and Two DBB

Fairs. The DBB Baseline begins Nine Sessions

after Surgery.

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