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Cortical Lesions and Sexual Behavior

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Gail Joan Sellstrom

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ABSTRACT
CORTICAL LESIONS AND SEXUAL BEHAVIOR IN THE MALE RAT

By

Gail Joan Sellstrom

The sexual behaviors of 39 naive adult male Long-
Evans rats were observed during three preoperative and
three postoperative 20 minute sessions. Between
preOperative and postoperative sessions 10 males
received full bilateral electrolytic lesions to the
cingulate cortex, 8 males received bilateral neocortical
lesions. and 9 males had sham operations. The remaining
12 males received no surgical treatment. Analyses of
fourteen measures of sexual behavior failed to yield
significant lesion group effects. Replication effects
were significant in three of the analyses. The results
confirmed the work of Beach (1940, 1941) and were
inconclusive with reSpect to the studies reporting
impaired ability to perform sequential responses

following cingulectomy.

CORTICAL LESIONS AND SEXUAL BEHAVIOR IN THE MALE RAT

By

Gail Joan Sellstrom

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Psychology

1973

\94 ACKNOWLEDGMENTS

I particularly wish to acknowledge the scholarly and
theoretical assistance provided by Professor Lawrence I.
O'Kelly. I wish to eXpress my gratitude to Professor
Raymond W. Frankmann who willingly advised with respect
to statistical aspects of the experiment. Dr. Glenn I.
Hatton who made available a great deal of technical
information as well as the surgical and histological
facilities necessary for the completion of the
eXperiment, and Dr. Lynn Clemens who suggested techniques
for manipulating reproductive cycles and provided
training in the observation and identification of sexual
responses. Computerized analysis techniques were made
available by Dr. F. Samuel Bauer. Also I wish to thank
Richard H. Radius for the constant encouragement so
patiently given during the final stages preceding

completion of this dissertation.

ii

TABLE OF CONTENTS

ACKNOWLEDGMENTS
LIST OF TABLES
LIST OF FIGURES
INTRODUCTION
METHOD

RESULTS
DISCUSSION
APPENDIX A
APPENDIX B
APPENDIX C

LIST OF REFERENCES

ii
iii

iv

28
36
69
75
77
103
106

LIST OF TABLES

Table l. Histological results for cingulate
and neocortical lesion groups. 37

Table 2. Results of unweighted-means two by
four factorial analyses of variance.
The harmonic mean is n.6602. 56

Table 3. Means and standard deviations for
preoperative and postoperative measures
of performance. 60

APPENDIX A. Table 9. Raw data: body weight
in grams. 75

APPENDIX B. Table 5. Raw data: autogenital
cleaning (a) and interresponse times
(irt) in seconds for mounts (m), mounts
with intromission (mi), and mounts with
ejaculation (me). 77

APPENDIX C. Table 6. Raw data: postejaculatory

responses not occurring within 20 minute
sessions. 103

iii

Figure 9. Mean difference score for preOperative
and postOperative mean interval between
ejaculations in replications I and II as a
function of treatment group.

Figure 10. Mean difference score for preoperative
and postoperative mean postejaculation
interval in replications I and II as a
function of treatment group.

Figure 11. Mean difference score for preoperative
and postoperative unadjusted mean number of
mounts per ejaculation in replications I and
II as a function of treatment group.

Figure 12. Mean difference score for preoperative
and postoperative adjusted mean number of
mounts per ejaculation in replications I and
II as a function of treatment group.

Figure 13. Mean difference score for preoperative
and postOperative unadjusted mean number of
mounts with intromission per ejaculation in
replications I and II as a function of
treatment group.

Figure 14. Mean difference score for preoperative
and postoperative adjusted mean number of
mounts with intromission per ejaculation in
replications I and II as a function of
treatment group.

50

51

52

53

Sh

55

INTRODUCTION

The influence of the cingulate cortex on reproductive
behavior in rats. hamsters. and cats has been investigated
by systematically destroying the cingulate cortex and
observing. under carefully controlled conditions. the
extent to which the preoperative behavioral sequence was
disrupted.

In 1940 and 1941 Beach reported that destruction of
20 percent of the cortex. regardless of site of
destruction, had no detrimental effect upon the sexual
behavior of male rats. Destruction of a greater
percentage, up to 80 percent, was found to result in
increasingly severe detrimental effects on the male rat's
behavior. Beach's results (1940. 1941) confirmed the
work reported by Davis (1939). Beach's (1940, 1941)
results, as represented in terms of Lashley diagrams
showing the percentage of cortex damaged, suggest that
Beach did not systematically destroy the cingulate
cortex. Also, it is not possible to determine how much
of the cortex was damaged since only the surface area
damaged and some vague descriptions of subcortical
damage are given. A more systematic approach to the
problem of sexual behavior and cortical destruction is

l

2

found in the reports of Larsson (1962. 1964). Larsson
(1964), on the basis of his work in 1962. concluded that
”bilateral lesions in the lateral parieto-temporal area

of the cerebral hemispheres have more severe effects upon
the mating behavior of male rats than injury to the medial
parts of the frontal. parietal, and occipital lobes.
Whereas lateral lesions permanently eliminated mating in
several of the animals and lowered the activity of others.
removal of the median cortex in no case did permanently
abolish sexual behavior.”

A survey of the literature suggests that eXperiments
dealing with the effects of cingulectomy upon behavior
patterns present in the animal's repertory at the time of
surgery and common to a species. i.e.. what are
sometimes called instinctive behavior patterns. have
involved the use of rats. hamsters, and cats. If one is
willing to grant that reSponse to emotional stimuli falls
within the general class of behaviors being considered.
i.e.. the instinctive. then the results of Bard and
Mountcastle (1948) suggest that the cingulate area may be
functionally related to the level of stimulus intensity
necessary to arouse characteristic responses. Bard and
Mountcastle (1948) reported that cats with cingulate
lesions show a decreased sensitivity to emotional
stimuli.

Food hoarding behavior following cingulectomy has

been investigated by Stamm (1954) and Bunnell and Pinder

3

(1964), the researchers using rats and hamsters,
respectively. A comparison of preoperative and
postoperative hoarding behavior revealed that rats with 8
to 17 percent of the median cortex damaged showed a large
decrease in the measure of hoarding behavior, these rats
also differing significantly from the relatively
unaffected control animals (lateral cortical lesions)
(Stamm, 1954). When groups were compared in terms of
latency of hoarding responses the differences were less
marked. Stamm (1954) also found evidence for a positive
relationship between severity of cingulate damage and
disruption in hoarding behavior.

According to Bunnell, 23 gl. (1966), Bunnell and
Pinder (1964) have demonstrated that Stamm's (1954)
results have some generality in that they have been
extended and confirmed for the hamster. Apparently
Bunnell and Pinder (1964) believe that "the lesion
effects were upon motivational factors, or, in terms of
Beach's theory (1955), upon the sexual arousal mechanism
(AM). The threshold of the AM, which is presumed to have
both a cortical and a subcortical component in mammals,
is influenced by (a) internal states and responses of the
organism, and (b) exteroceptive stimulation" (Bunnell.
gt gl., 1966). In an investigation of cingulate effects
on duration, amount, and kinds of sexual responses in the
male syrian golden hamster, Bunnell, gt g1. (1966) report-

ed that cingulate postoperative behavior differed

LL

temporarily from preoperative behavior in respect to
number of intromissions made, duration, and the ability to
maintain the typical position during autogenital cleaning,
the frequency of intromissions being significantly
decreased. the time required to perform standard
responses being longer, and some imbalance during auto-
genital cleaning being evident. The only differences
observed by the end of postoperative testing involved
frequency of intromissions. recovery being incomplete in
four of the seven cingulectomized hamsters. There was no
immediately apparent disruption in motor behavior
(Bunnell. gt_§l.. 1966). Although extracingulate areas
damaged included the dorsal hippocampus, corpus callosum,
fornix, and dorsal septum, Bunnell, 33 El. (1966) found
no evidence for a relationship between extent of lesion
and behavior of cingulectomized subjects.

The results obtained by Stamm (1955) and Slotnick
(1967), in their investigations of the effects of
cingulectomy on maternal behavior in rats, parallel those
reported by Stamm (1954) and Bunnell and Pinder (1964)
with respect to hoarding behavior. The experiments of
Slotnick and Stamm differed in that Stamm's (1955) rats
had delivered many litters prior to serving as subjects
while Slotnick's (1967) rats were primiparous at the time
of surgery. Both investigators reported that following
cingulectomy maternal behavior, including such responses

as nest building, retrieving of pups, and string pulling,

5

was severely disrupted. Thus, the number of litters
delivered prior to surgery does not appear to be an
important variable with reSpect to the effects of
cingulectomy on maternal behavior.

General theoretical accounts of the functional
significance of the cingulate area are summarized by
Bunnell, gt a1. (1966) as follows: MacLean (1958)
favors the notion that the limbic system is implicated in
the control of instinctive behavior patterns, there being
evidence “that certain limbic structures, including
median (cingulate and retrosplenial) cortex, are
particularly important to behaviors promoting survival of
the species (e.g.. reproductive activities)." As noted
by Bunnell. gt El. (1966), Pribram has suggested that the
”essential function of the limbic system is the
integration of behavioral components into smoothly
functioning sequences and...the changes in instinctive,
affective, or other classes of behavior which follow
limbic manipulations are the result of alteration or
disruption of the sequencing of acts which comprise such
complex behaviors.” With the exception of the eXperiments
by Beach (1940, 1941), Davis (1939), and Larsson (1962),
the evidence from experiments involving hoarding,
maternal, and sexual behavior has tended to provide
support for Pribram's theory.

Acquisition and retention following cingulectomy

have been the focus of interest in studies in which

6

various measures of learned behavior are viewed as the
dependent variables and attempts are made to demonstrate
functional relationships between these performance
measures and various manipulations in the cingulate
region. These eXperiments may be distinguished by
appealing to task characteristics. Thus, studies
involving some form of noxious stimulation which the
animal escapes or avoids by engaging in a Specific
activity include running to escape shock in a T-maze,
bar-pressing to terminate a loud noise, withholding a
reSponse (e.g.. drinking) to avoid shock (passive
avoidance), and performing a Specific response (e.g..
running to the opposite side of a shuttle-box when the CS
comes on) to avoid shock (active avoidance).

Before considering the first loosely related set of
acquisition and retention studies it might be of value to
discuss briefly the major issue involved. In most
instances cingulectomy results in disruption of original
learning of active avoidance responses and has no
apparent effect on original learning of passive
avoidance reSponses. Two major accounts of the
functional significance of the cingulate cortex have been
preposed in response to these results. Lubar and
Perachio (1965) distinguish between the response-
facilitation and the fear-facilitation drive hypotheses.
Central to the reSponse-facilitation hypothesis is the

notion that the cingulate area facilitates behavioral

7

responsivity and that removal of the cingulate area might,
therefore. be eXpected to result in an opposite pattern of
effects, namely lessened behavioral responsivity. The
fear-facilitation dirve hypothesis consists of the notion
that this fear facilitation may be significantly related
to inferior active avoidance reaponding. Two findings
relating to the latter theory are those of Bard and
Mountcastle (1950), in which cingulectomy reduced
sensitivity to emotional stimuli, and Trafton (1965), in
which cingulectomy increased freezing in reSponse to what
was presumably a fear stimulus.

The following three studies may be viewed as
satisfying two cirteria: a) they involve some form of
noxious stimulation: and b) they do not involve passive or
active avoidance responding. Brady and Nauta (1953)
observed CER retention in rats following training and
surgical procedures appropriate to membership in the
septal. cingulate, or unaperated control group. Brady
and Nauta (1953) reported that only the septals showed
greater emotional reactivity and an increase in the
magnitude of the startle response. An experiment
involving the effects of combined limbic, adjacent
cingulum (area cingularis anterior ventralis of LA). and
striae lesions on retention of lever-pressing-to-
terminate-a-noxious stimulus behavior, was conducted by
Lyon and Harrison (1959). In terms of extent of

disruption of lever pressing to terminate a 105 db noise

8

for 20 seconds. the complete cingulate and control rats
were very similar, further evidence suggesting that
partial destruction of the area cingularis anterior
ventralis results in no disruption of the lever pressing
response (Lyon and Harrison. 1959). In terms of amount of
time required to attain preoperative performance levels.
Lyon and Harrison (1959) found that the distributions of
experimental and control groups showed a considerable
amount of overlap.

While the two studies just cited were primarily
concerned with cingulate effects on memory or previously
learned behavior the study of Thompson and Langer (1963)
was focused upon original learning in rats. After their
animals had learned to run to escape shock in a T-maze
Thompson and Langer (1963) subjected the rats to lesions
at several sites (summarized with accompanying results
below) and, 2 weeks later, tested the animals on a
reversal of position task. In terms of the effects of
lesions on reversal performance the eight groups were.
described by Thompson and Langer (1963) as follows: (a)
normal and neocortical controls showed no effect, i.e.. a
high level of accuracy: (b) precallosal anterior limbic,
hippocampal, septal, and preoptic hypothalamic animals
were significantly inferior to controls: and (c)
supracallosal anterior limbic and fornix column animals
were not significantly deficient. The authors also

reported that reversal performance was not effected by

9

lesions in the pretectal or parafascicular area of the
thalamus, the subthalamus, the substantia nigra, or the
amygdala (Thompson and Langer. 1963).

Several investigators have been concerned with
cingulate lesions and passive avoidance responding.
Kaada, 33 El. (1962), working with rats, and McCleary
(1961), Lubar (1964), and Cornwell (1966), all working
with cats having mid-cingulate lesions, have reported that
cingulectomized animals do not differ from normal animals
in acquiring a passive avoidance response. Lubar (1964)
found that cats with combined limbic cortex-septal area
and cingulate lesions showed normal passive avoidance
acquisition, while animals with damage limited to the
limbic cortex-septal area were inferior to normals with
respect to passive avoidance acquisition. Lubar's (1964)
cats with lesions limited to the mid-cingulate area were
more resistant to extinction of the passive avoidance
response than were normal cats or cats with combined
lesions. Kaada (1962) also reported that passive
avoidance acquisition was normal regardless of what parts
of the cingulate cortex and adjacent corpus callosum were
damaged. i.e.. the medial cortex lying in front of and
above the corpus callosum, or the posterior cingulate
cortex and/or retrosplenial cortex.

Before discussing typical active avoidance
eXperiments it might be noted that a study conducted by
Pribram and Weiskrantz (1957) differs from the most

lO

widely cited studies in terms of subjects. design, and
results. Rhesus monkeys were trained to avoid shock in a
shuttle-box prior to surgery, and, following a one week
recovery interval, were tested successively under
extinction, reacquisition, and extinction conditions.
Although active avoidance behavior was observed following
lesions in several areas other than the medial frontal and
cingulate cortex the only directly pertinent results were
that cingulates did not differ from controls in terms of
either extinction 1 or reacquisition performance and were
superior to the controls (i.e.. reached the extinction
criterion more rapidly) in terms of extinction 2
performance (Pribram and Weiskrantz, 1957).

With the exception of the Peretz (1960) eXperiment,
all of the studies below, focusing upon the acquisition
of an active avoidance response following cingulectomy in
rats, involved the use of a two-way shuttle box. When
performance was evaluated in terms of trials to criterion,
or a related measure, Thomas and Otis (1958), Thomas and
Slotnick (1963), Trafton (1965), and Peretz (1960) found
that cingulectomy resulted in inferior performance.

One might note that in addition to damaging cingulum
fibers connected to the posterior cingulate cortex, the
eXperimental animals of Thomas and Otis (1958) sustained
bilateral hippocampal damage. In the two eXperiments
conducted by Thomas and Slotnick (1962), in which rats

were postOperatively exposed either to CAR training

11

followed by maze training (Exp. 1) or to the two
procedures in reverse order (Exp. II), it was also found
that cingulate lesions having an observable effect in the
shuttle-box situation have no effect in the maze situation,
regardless of procedure order, whereas prior maze learning
significantly improved later CAR performance, the latter
result being attributed to the handling of animals
occurring during maze training.

The handling effect noted by Thomas and Slotnick
(1962) might be considered in evaluating McCleary's (1966)
account of Peretz's 1960 results. As noted earlier,
Peretz (1960) obtained the typical active avoidance
acquisition result in spite of his use of a one-way
procedure in which rats avoided shock by running from a
black to a white compartment. McCleary (1966), on the
basis of the Candland, 33 gl. (1962) finding that rats
will learn to avoid a black compartment when the
consequence of being present in the black compartment is
a 20 second period of being stroked, head to tail, while
being held in a gloved hand, claims that handling is
aversive to the rat. McCleary (1966) further speculates
that Peretz (1960) ”may have introduced an approach-
avoidance conflict into his one-way procedure" which
”augments the avoidance conflict in cingulectomized rats"
to the point where the deficit is “great enough to be
manifested in the one-way situation.” Before continuing

it might be of value to note that Peretz's (1960)

12

cingulate animals were also inferior to shams, in terms of
both total trials and oscillation trials to criterion,
when oscillation was substituted for shock as the aversive
stimulus. In terms of resistance to extinction,
cingulates took more trials to criterion under oscillation
conditions and did not differ from shams under shock
conditions (Peretz, 1960).

Thomas and Slotnick (1963), in addition to finding
the characteristic deficit under 2 to 3 hours food
deprivation conditions, found that cingulectomized rats
fail to show the deficit in CAR performance when run
under high drive (daily feeding occurs immediately after
trials) conditions. Thomas and Slotnick (1963) account
for their results by suggesting that ”lesions affect
performance by enhancing the tendency of rats to freeze in
response to the CS and the high hunger drive counteracts
the freezing response by inducing heightened general
activity which protects §§ from lesion-induced behavioral
loss.” Evidence regarding differential effects of full,
anterior, or posterior cingulate damage was provided by
Trafton (1965). Either full or anterior cingulate lesions
led to a very severe deficit: i.e.. 24 of the 25 rats
comprising these two groups failed to show any evidence of
learning the shuttle-box avoidance response within 100
trials. while rats with posterior cingulate lesions tended
to freeze when running was apprOpriate and thereby showed

a deficit in CAR responding.

13

Although the results for cats are less consistent it
is not clear that cats differ markedly, if at all, from
rats in respect to acquisition of an active avoidance
reSponse following cingulectomy. For example, results
obtained by Cornwell (1966) are regarded as evidence that
mid-cingulate lesions disrupt acquisition of an active
avoidance response. However, Lubar (1964) reports that
the mean trials to criterion for all four groups
(combined limbic cortex-septal area and cingulate gyrus,
limbic cortex-septal area. cingulate gyrus, and normal
controls) were nearly identical, and later expresses
suprise at failing to confirm McCleary's (1961) finding
with cats. Cornwell (1966) found that cingulectomized
cats required a mean of 67.7 (range of 25 to 200) trials
to the active avoidance criterion while sham operated
cats required a mean of only 39.5 trials (range of 14 to
70). Cornwell's cingulates also required more first
retraining and second retraining trials than shams, the
two groups being very similar in terms of crouching to
the CS and measures of urination and defecation. McCleary
(1966) suggests that the discrepancy between his own
results (1961), as well as those of Cornwell (1966), and
those of Lubar (1964) could be accounted for if one were
willing to view the one-way avoidance task used by
Lubar (1964) as easier than the two-way task used by the
other two investigators.

In 1964 Moore exposed cats to CAR training

14

procedures in a double-grill shuttle-box prior to removal
of the cingulate cortex and subsequently measured
retention during no shock trials. In addition to finding
that 5 out of 6 cingulates failed the retention criterion,
Moore (1964) also found that, relative to septal animals
and animals with combined (septum and hippocampus)
lesions, cingulate animals were least effected in terms
of retention and relearning, the 5 cingulate animals
failing the retention test subsequently relearning in
fewer trials than were required to reach the criterion
preoperatively. 0n the basis of these results Moore
(1964) postulates that while the cingulate cortex may
play a role in habit retention, this role is not
essential for retention or relearning.

A further confirmation of McCleary's (1961) findings
is provided in the report of Lubar and Perachio (1965).
These authors, in addition to demonstrating that
cingulate cats are clearly inferior to controls with
respect to two-way active avoidance acquisition and
somewhat inferior to controls with respect to one-way
avoidance acquisition, found that both controls and
cingulates receiving one-way training were clearly
superior to comparable groups receiving two-way training
(Lubar and Perachio, 1965). Following acquisition each
group was then given training in the alternate situation,
the following series of results being obtained for the

transfer sessions: a) regardless of group membership

15

animals transferred from one-way to two-way avoidance
were inferior to animals transferred from two-way to one—
way avoidance training: b) cingulates were similar to
controls during one-way transfer and clearly inferior to
controls during two-way transfer: and c) while control
animals transferred to two-way avoidance were superior to
animals originally receiving two-way training, cingulates
did not differ from controls with respect to this measure
(Lubar and Perachio, 1965). Lubar and Perachio (1965)
also reported that during transfer training on the two-
way task subjects, regardless of group membership,
vocalized more frequently than subjects transferred to
the one-way task. The authors felt that their results
supported the fear-facilitation drive hypothesis, their
conclusion being that fear in cats may be facilitated by
cingulate lesions and that this facilitation may be a
significant determiner of the active avoidance response
deficit (Lubar and Perachio, 1965).

The discussion will now focus upon behavioral studies
in which acquisition or retention of an instrumental
response takes place without aversive stimuli. The scepe
of this discussion will be further limited by excluding
experiments involving lever pressing behavior.
Experiments satisfying these criteria involve visual
discrimination. funnel displacement, and leg displacement.
Because of their immediate relevance with respect to the

present experiment, the alternation studies will be the

16

last non-aversive instrumental response studies
considered.

Pribram, gt_§l. (1962) protested rhesus monkeys in a
modified Wisconsin General Testing Apparatus, subjected
them to surgical procedures designed to destroy pre-,
sub-, and supracallosal cortex, and then gave them visual
discrimination training. Pribram, g3 El. (1962) found
that cingulates were slightly inferior to normals in terms
of trials and errors to criterion. Peretz (1960), in the
series of experiments previously cited, trained hungry
cingulate and sham operated rats to discriminate between
black and white cues. Using the correction procedure.
Peretz (1960) found that cingulectomized animals were
significantly superior to sham operated animals in terms
of both trials and number or errors to criterion. there
being no differences between groups in terms of latency
of response on criterional trials. The cats used in the
avoidance study of Cornwell (1966) learned to displace a
funnel for food according to a VR schedule. Cingulates
required a mean of 5.3 sessions (range of 2 to 10) and
shame required a mean of 4.7 sessions (range of 0 to 10)
to reach the criterion of having at most a 5 second
latency on at least 90 of the 100 daily trials for 3
consecutive days. Cornwell's (1966) cingulectomized cats
were also similar to the sham operates during extinction
of the funnel displacing response.

A further eXperiment falling within the category of

17

studies involving difficult to classify tasks was
conducted by Brutkowski and Mempel (1961). Prior to
destruction of either the rostral cingulate or the
posterior cingulate area dogs were trained to reSpond
differentially on the basis of a discrimination between
two tones reflecting 08+ and 08-, food reinforcement
being presented only when the foreleg was placed on the
food tray during CS+ presentation. Errors included
failing to place the foreleg on the food tray during CS+
presentation and placing the foreleg on the tray during
08-. The authors obtained evidence suggesting that lesion
site is related to the extent of instrumental response
retention in that posterior cingulectomy failed to disrupt
retention performance while rostral (genual area) damage
resulted in a marked failure to inhibit responding during
03-, 4 to 15 days of relearning being required to regain
preoperative performance levels. Brutkowski and Mempel
(1961) felt that their rostral cingulates made more
reward motivated reSponses, expected food regardless of
which CS was being presented, and were more vigorous in
taking the food. These results thus confirmed the
earlier work of Brutkowski and his associates in which
genual (or rostral) cingulectomy resulted in a temporary
inability to withhold defensive CR3 and an increase in
correct and incorrect responses, emotional reSponses
(violent rage and angry behavior), and fear-like

responses. These results are regarded as evidence

18

suggesting that ”the genual portion of the anterior
cingulate area is one of the critical forebrain regions
for the inhibition of some affective reSponses”
(Brutkowski and Mempel, 1961).

It has been found that cingulectomy may either
markedly facilitate (Peretz, 1960) or mildly disrupt
(Pribram, g1 g1., 1962) acquisition of behavior patterns
involving visual discrimination. Cornwell's (1966)
study, lacking the discrimination element common to the
other three studies, has yielded results easily falling
within what appears to be the normal range: i.e..
Cornwell's (1966) cingulectomized cats were similar to
Pribram, gt gl;§ (1962) cingulectomized monkeys in being
only slightly, and adversely, affected. Continuing this
negative trend, it has been found that cingulectomy not
only leads to marked retention deficits but also leads to
a marked increase in emotional responses (Brutkowski and
Mempel. 1961). This latter result might be regarded as
disconfirming, in some weak sense, the earlier results
of Bard and Mountcastle (1950).

Attention shall now be focused upon experimental
attempts to evaluate the effects of cingulate lesions on
various aspects of lever pressing behavior. 0f the seven
bar pressing eXperiments to be considered only three
involve the retention design while five represent
attempts to assess the effects of lesions on original

learning. The latter set includes three eXperiments in

19

which reinforcement is provided according to DRL schedules,
those of Stamm (1963, 1964) involving rhesus monkeys
while Ellen, Wilson, and Powell (1964) worked with rats.

Using delays ranging from 10 to 70 seconds, Stamm
(1963) found that cingulates were similar to normals for
delays up to and including 30 seconds, no cingulate
meeting the acquisition criterion when longer delays were
used. Normal monkeys were able to meet the learning
criterion when delay intervals of less than 60 seconds
were in effect (Stamm, 1963). In his 1964 study Stamm
tested each animal by making the delay interval 60
seconds longer than the last delay interval experienced
during training and found that cingulate monkeys were
superior to normal monkeys in terms of rates of multiple
presses: i.e.. significantly higher rates of multiple
responses (responses occurring within 2 second intervals)
were observed for normal monkeys. Stamm (1964) also
analyzed interresponse time distributions and reported
that the timing responses of normals were less clear
than those of cingulate animals. Stamm (1964) speculated
that the multiple presses of the normal monkeys might
reflect frustrative behavior while the superiority of the
cingulates might be related to motivational functions of
the cingulate cortex. The results reported by Ellen,
Wilson. and Powell (1964), using rats and a 20 second
DRL schedule, were comparable to those reported by Stamm

(1963) for monkeys: i.e.. cingulectomized rats acquired

20

the timing response as readily as normal rats.

As the final eXperiment in his series of four,
involving the same subjects throughout the series such
that the rats were lesioned 8 months prior to the
eXperiment now being discussed, Peretz (1960) trained his
cingulectomized and sham Operated rats to bar press for
food according to a CRF schedule. The animals were
maintained at a level 10 to 15 percent below normal body
weight. Following CRF acquisition Peretz (1960)
substituted a VI schedule (range of 10 seconds to 7
minutes: mean of 3 minutes) for the CRF schedule, the six
days using the VI schedule being regarded as a test
sequence. In terms of both rate of bar pressing. i.e..
number of responses per 30 minute session, and mean rate
over the six sessions, the cingulates were significantly
superior to the sham Operates (Peretz, 1960). It is
interesting that in two eXperiments using subjects
representing quite distinct levels of evolutionary
development and involving quite distinct temporal
reinforcement schedules the results are comparable to the
extent that cingulectomy facilitates original learning of
the lever pressing response (Stamm, 1964: Peretz, 1960).

In one of the few studies involving water reinforced
operant response learning Ellen and Powell (1962)
compared the acquisition performance of septal and
cingulate lesioned rats under a multiple reinforcement

schedule comprised of FR 15 and PI 2 minute schedules.

21

Although Ellen and Powell (1962) did not find that
acquisition performance varied as a function of lesion
site they were able to report that the rate of responding
by cingulates under the VI 2 minute schedule was
significantly inferior to the corresponding behavior of
the septal animals.

The eXperiment of Ellen and Powell (1962) was
designed so as to permit the collection of data relevant
to the evaluation of the effects of cingulectomy on
retention performance. For those animals trained on
PR 15 and VI 2 minute schedules prior to surgery
retention testing was carried out on postoperative days
one through twelve. Retention performance under FR 15
conditions did not differ for either group from
preoperative performance. However, Ellen and Powell
(1962) did find that retention performance under the VI 2
minute schedule was related to site of lesion, septals
showing a marked and permanent increase in number of
responses emitted while cingulates showed only a
temporary increase in the number of responses made during
the final part of the interval and continued to pause
following reinforcement.

Extending the work of Stamm (1963). Glicketein. 31
El. (1964) trained monkeys to respond according to a DRL
reinforcement schedule, subjected them to frontal lesions
similar in extent to those reported by Stamm (1963), and

then conducted postoperative retention tests. The

22

pattern of results presented in these two reports might
be regarded as a reversal of the usual pattern: i.e..
Glickstein, §t_§l. (1964) found that cingulate lesions led
to a disruption in retention of DRL performance whereas
Stamm (1963) reported that cingulate lesions failed to
influence DRL acquisition. Because Stamm's (1963)
training procedure involved the presentation of what
could be regarded as a consistent discriminative stimulus
(Z-second white light after each reinforcement).
Glickstein, 33 El. (1964) feel that the animals were
being trained "not to respond in the presence of a light“
in the first 2-second interval, ”a procedure which limits
any conclusion about timing per se.“

The third and final bar press retention study to be
considered involved the preoperative training of rats to
bar press in the presence of a discriminative stimulus.
food being provided as reinforcement in accordance with a
VI 15 second schedule, and the postOperative testing of
retention and extinction performance (Schwartzbaum, g1
,§1.. 1964). Schwartzbaum, £3.31. (1964) found that.
relative to preoperative performance, cingulate
responding during presentation of the reinforced
discriminative stimulus (3000 cps pulsing tone at 70 db)
was greatly reduced while responding during the presence
of the non-reinforced stimulus (550 cps tone at 70 db)
was not altered. The authors also found that extinction

was facilitated by cingulectomy: i.e.. the cingulates

23
showed less resistance to extinction than sham operates
(Schwartzbaum, 35 gl.. 1964). The extinction results are
accounted for in terms of the assumed function of the
septal area and McCleary's (1961) concept of ”response
specificity" which assumes that ”the facilitatory and
inhibitory systems which control response tendencies
normally operate in some reciprocal relationship to one
another,” such that “damage to one would increase the
effects of the other....” (Schwartzbaum, g3 g1., 1964).
That is, removal of the cingulate area increases outflow
from the inhibitory system (intact septal area) and
thereby results in a relatively rapid cessation of
responding. In commenting on the somewhat anomalous
results reported by Peretz (1960), Schwartzbaum, 33 gl.
(1964) speculate that the superior visual discrimination
learning shown by cingulectomized rats may be attributable
to ”enhanced inhibition of incorrect response tendencies
...” and later observe that the superior VIZ minute lever
pressing performance of cingulate subjects reported by
Peretz (1960) ”was not evident in the cingulectomized
subjects in the present study.”

The alternation studies to be considered involve a
design in which training is followed by surgery and
effects of brain damage are evaluated by comparing
postOperative and preoperative measures of performance
on the same subjects as well as postoperative and

preoperative measures between subjects. The results

24

provided by such experiments might be regarded as
relevant to the question of the extent to which the
cingulate area is an important determiner of processes
related to original learning, retention of original
learning (or memory), both of these, or, neither of these.
The alternation studies also provide results relevant to
Pribram's (1966) hypothesis involving the limbic system
and the integration of responses in a given sequential
task. It might be noted that Gross, g1 51. (1965) found
that a deficit in original learning as well as in the
retention of alternation behavior may result from lesions
of the caudate nucleus, of the anterior cortex, of the
hippocampus, or of dorsal thalamic structures. Thus the
evidence does not appear to support the notion that the
cingulate cortex has a unique function in the relevant
processes.

The results reported for rhesus monkeys by Pribram,

l. (1962) and Pribram, 33 Q1. (1966) suggest that

g3
retention of neither a delayed alternation task nor of
right left and go-no-go alternation is disrupted as a
result of cingulate damage. However, in the former
experiment acquisition of the delayed alternation task
was disrupted following cingulectomy (Pribram, £1 21..
1962). In both cases an attempt was made to include the
projection sector of the anterior thalamic nuclear group
in the region destroyed (Pribram, g3 g1.. 1962: Pribram,

a: 11:09 1966).

25

The retention eXperiments cited seem to share a
characteristic, namely, measures of retention performance
do not appear to discriminate between normal and
cingulectomized animals. Fortunately it is still
reasonable to doubt the generality of this finding.

Using a complex measure of behavior, consisting of the
starting and running speed ratios of nonreinforced to
reinforced trials. Barker and Thomas (1965) found that
full cingulate lesions led to a significant disruption of
acquisition and retention of a runway alternation task.
Barker and Thomas (1965) also found that only one of five
cingulate rats reached the relearning criterion within
the 200 trials permitted, the remaining four rats failing
to show any indication of retention or relearning.

0n the basis of evidence obtained from eXperiments
involving Species-specific behaviors of the kind
discussed in the present eXperiment, as well as the
evidence reported by Michal (1965), Thomas, Hostetter,
and Barker (1968) suggest that ”the effects of lesions in
dorsal limbic cortex on species-specific maternal and
sexual behavior have indicated that mechanisms of
temporal-response integration were impaired.” The
results of the series of studies conducted by Barker
(1965) and Barker and Thomas (1965, 1966) are consistent
with the hypothesis that the impaired functioning of
mechanisms of temporal-response integration ”might be

evident in a behavioral end point in which a learned

26

sequence of responses was the dominant feature" (Thomas,
HoStetter, and Barker, 1968).

The present experiment was conducted in order to
determine the extent to which the cingulate cortex is, or
is not, necessary to typically observed sexual behavior
in the adult male Long-Evans rat. Preoperative and
Postoperative sexual behavior was measured in terms of
latency to first mount without penetration (mount),
latency to first mount with brief penetration
(intromission), latency to first mount with penetration
and ejaculation (ejaculation), inter-reSponse interval,
number of mounts and intromissions occurring prior to
each ejaculation. frequency of mounts, frequency of
intromissions, frequency of ejaculations, postejaculatory
interval. and presence or absence of autogenital cleaning
between intromissions. These measures of sexual behavior
were selected from those described in the reports of
Bermant, 33 Q1. (1968), Dewsbury (1967), Beach (1956),
Beach and Jordan (1956), and Beach and Whalen (1959).

On the basis of the evidence reviewed, the
exceptions including the reports of Beach (1940, 1941),
Davis (1939), and, perhaps Larsson (1962), it was
supposed that full cingulectomy would lead to a
disruption in the typical behavior pattern exhibited by
the adult male Long-Evans rat. More specifically, it was
supposed that the experiment would provide evidence

indicating a failure to complete sequences of behavior

27

begun, such evidence being comparable to and consistent
with the results obtained in studies of maternal behavior
(Slotnick, 1967) and studies of alternation learning
(Barker and Thomas, 1965, 1966).

This supposition was
not confirmed.

METHOD

Thirty-nine naive male and thirty naive female rats
of the Long-Evans strain were obtained from the Chordata
Corporation of Ontario, New York. The animals were 80 to
90 days old when first received and exposed to pre-
experimental conditions. The subjects were 94 to 104 days
old when preoperative testing began. The subjects were
130 to 140 days old at the termination of the experiment.
The males served as eXperimental subjects while the
females were used as stimulus animals. The males were
housed one per cage whereas the females were housed six
per cage, cages for males being 20.9 cm. long, 15.4 cm.
high, and 17.6 cm. wide. Cages for females were 20.9 cm.
long, 15.4 cm. high, and 57.2 cm wide. All cages were of
the wire bottomed variety manufactured by the Wahmann
Company. Water and Wayne Mouse Breeder Blox were
available gg‘11b in the home cages throughout the
experiment. All eXperimental subjects were weighed every
other day prior to surgery. Following convalescence the
subjects were again weighed every other day. When it
became necessary to terminate one of the experimental
subjects due to pneumonia, all animals involved in the

experiment were given an intramuscular injection of

28

<wf‘

t

29
penicillin. The injections were administered on the last
day of surgery, between the last testing session for that
day and the start of surgical procedures.

A reversed light-dark cycle was used in order to
maximize the probability of observing subjects during
their periods of greatest sexual responsiveness.
Replication I animals were housed in the experimental
room and subjected to observation during the dark portion
of the light cycle. Because of their unanticipated early
arrival, replication II subjects were housed in a
similar but separate room during the first 11 days of
the reversed light-dark cycle. Replication 11 animals
were moved to the eXperimental room three days prior to
session one. Replication II subjects were then maintained
in the eXperimental room until termination of the
experiment.

All animals were exposed to the reversed light cycle
for 14 days prior to being observed under eXperimental
conditions. Two Knight all purpose automatic reset
timers were used to control the light cycle. After 10
hours of darkness a 100 watt lamp was turned on by one of
the timers, this lamp remaining on for 14 hours. One
hour after the first 100 watt lamp was turned on a second
100 watt lamp was turned on, this lamp remaining on for
12 hours. The dark part of the cycle was in effect
during the time interval beginning at 7 a.m. and ending

at 5 p.m.. all observation procedures being carried out

30
during these hours. Injections were administered during
the latter part of the light portion of the light cycle.

All stimulus females used during preoperative and
postoperative testing were brought into estrous once
every 5 days. Ninety-six hours prior to the observation
session the females were given a .05 cu. cm. subcutaneous
injection of l microgram per .05 cu. cm. estradiol
benzoate in sesame oil. This injection procedure was
repeated 72 and 48 hours prior to observation. Six hours
prior to observation the same females were given a .1
cu. cm. subcutaneous injection of .5 mg. per .1 cu. cm.
progesterone suspended in sesame oil.

The apparatus consisted of a rectangular five sided
plexiglass observation box and Esterline Angus recording
equipment. The observation box was 50.6 cm. long. 37.4
cm. wide, and 37.4 cm. high, the missing panel in the
rectangular box being the top panel. The plexiglass
panels were approximately .5 cm. thick. During the
observation sessions the floor of the observation box
was covered with Royal Craft Cobmeal to a depth of
approximately 6 cm. Indirect light was provided by a 15
watt lamp and animals were observed in an area screened
off from the living cages.

Recording equipment consisted of a 20 channel
Esterline Angus Recorder and a remote control panel,
both of which operated through a 110 volt source. Two

channels of the recorder were used and a paper speed of

31

3.8 cm. per minute was used throughout the testing
sessions. During each observation session E sat
approximately one meter from the front panel of the
observation box. The recorder remained on during any
given session, the beginning and end of any given
observation period being indicated by pressing one of the
two buttons mounted on the remote control panel and
connected to the terminals of the recorder. One channel
of the recorder was used to indicate the limits of the
observation period. mounts without intromission, and
sexual behavior followed by autogenital cleaning. The
second channel was used to indicate mounts with
intromission and ejaculations,the latter initialed on the
recording paper by E as "E."

During preoperative and postOperative testing all
animals were systematically exposed to the observation
apparatus and members of the opposite sex. A copulating
male, not used as an experimental subject, was used to
test each female for receptivity. Only females exhibiting
the lordosis response were used during the test sessions.

Three 20 minute preoperative testing sessions were
carried out using females which had met the receptivity
criterion, i.e.. made at least one lordosis response with
a non-eXperimental male. The recording apparatus was
operating when the animals were placed in the observation
box. The male was placed in the observation box for 10

minutes prior to the beginning of any test session. After

32

the male had spent 10 minutes in the box E placed a
stimulus female in the center of the box, one pen on the
recorder being deflected as the female was placed in the
box. Any sexual behavior occurring during the 20 minute
test session was recorded by deflecting one of the two
pens on the Esterline Angus Recorder. A final deflection
of the first pen occurred at the end of the 20 minute
session. Each male was returned to the home cage
following any given test session. Only males which had
ejaculated at least once during preOperative session one
were used as experimental subjects. Preoperative sessions
two and three were conducted using the procedures
described for session one. Two or three days intervened
between sessions one, two, and three.

0f the 42 males observed during sessions one, two,
and three, 10 were randomly assigned to the cingulate
group, 10 were randomly assigned to the neocortical
group, 10 were randomly assigned to the sham group, and
12 were randomly assigned to the normal group. Five
cingulate subjects appeared in each replication. Due to
illness, only 3 neocorticals appeared in replication I.
Replication II involved 5 neocortical subjects. Sickness
accounts for the fact that only 4 shams appeared in
replication I. Replication II involved 5 sham subjects.
Six normal subjects appeared in each replication.

All surgery was performed under ether anesthesia.

Following anesthetization and head shaving, each animal

33

was immobilized by ear bars and bite bar of a Stoelting
Stellar stereotaxic instrument. In order to avoid
puncturing the ear drums, small bits of cotten were
placed in the ear canals of each rat prior to placing the
rat in the stereotaxic instrument. The top of the skull
was exposed and the stereotaxic instrument was used to
locate the point at which holes were to be drilled and to
guide the electrode to the appropriate depth. The
electrode consisted of a four cm. length of stainless
steel dental wire .25 mm. in diameter, the electrode
being insulated with epoxylite except for a .5 mm. tip.

In order to prepare the animal for cingulate lesions
10 small holes were drilled 0.8 mm. lateral to the
midline on each side of the skull. The holes were drilled
1.0 mm. apart and extended from a point 4.5 mm. anterior
to bregma to a point 4.5 mm. posterior to bregma. Full
cingulate lesions were made at depths of 2.0 to 4.5 mm.
below the skull in accordance with a brain map provided
by David J. Barker (1966) and subsequently modified by E.
In the case of each neocortical animal 10 small holes
were drilled 1.5 mm. lateral to the midline on each side
of the skull, the holes being spaced 1.0 mm. apart and
being located symmetrically with respect to direction
away from (posterior to or anterior to) bregma. All
neocortical lesions were made at a depth of 2 mm.
beneath the skull. In both lesion groups, i.e..

full cingulate and neocortical, the dura was punctured

34

and anodal electrolytic lesions were produced by passing
a 1.5 ma. direct current through the uninsulated tip of
the electrode for 10 seconds. The lesion producing
device was manufactured by the Stoelting Company.

Sham operation animals were subjected to the same
surgical procedures used with cingulate and neocortical
animals with the exception that Operations were completed
without puncturing the dura or in any way damaging the
cortex. Normal animals were subjected to no surgical
procedures.

The first postoperative tests were conducted 10 days
after the completion Of surgery. Stimulus females were
brought into estrous and selected for receptivity as
described for the preOperative tests. The three
successive 20 minute postoperative test sessions were
conducted in accordance with the procedures established
prior to and existing during the preoperative testing
sessions.

Upon completion of the third postoperative testing
session the animals were sacrificed by overeXposure to
ether and perfused with physiological saline followed by
an approximately 10 percent formalin solution. The
brains were removed from the skulls immediately after
perfusion and stored in approximately 10 percent formalin
for at least three days. The brains were then transferred
to a sucrose and formalin solution for at least three days

prior to slicing and mounting. The brains of all

35

cingulate and neocortical animals were frozed and sliced
into sagittal sections 50 microns in thickness, every
section being mounted and stained. A cresyl violet Nissl
stain for cell bodies was used. Using a microprojector
and a magnification of 15X, alternate sections were
projected for sketching. The extent Of cortical damage
was evaluated by measuring each sketch with a compensating

polar planimeter.

RESULTS

Alternate sagittal sections were examined for each
cingulate and neocortical animal completing the
eXperiment. The results of histological analysis are
summarized in Table 1. Total tissue damaged in cingulate
animals varied from 24.5 to 61.5 cu. mm. Gliosis
accounted for 6.5 to 26.5 cu. mm. of damage in the
cingulate group, while 17.0 to 36.5 cu. mm. of brain
tissue were completely destroyed in this group of
animals.

Due to severe subcortical brain damage caused by a
blood clot detected during histological analysis, data
from neocortical subject number 20 (total damage involved
139.0 cu. mm. of brain tissue) are not included in any
of the analyses.

Total brain tissue damage in neocortical subjects
ranged from 26.0 to 90.0 cu. mm. Gliosis was apparent in
9.5 to 36.0 cu. mm. of brain tissue in neocortical
animals, 16.5 to 57.5 cu. mm. Of completely destroyed
tissue accounting for the reamining damage.

Evaluation of pilot lesioned brains in terms of
sketches on Lashley diagrams indicated that 10 percent or

less Of the surface area Of the cortex is destroyed when

36

37

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41

lesions damage 24.4 to 39.0 cu. mm. of brain tissue.

In addition to severe cingulate damage, cingulates
showed damage to a minor degree to the corpus callosum,
dorsal hippocampus, frontal pole, and the anterior
Olfactory nucleus. Neocortical subjects showed extensive
neocortical damage as well as minor damage to the dorsal
hippocampus, corpus callosum, and caudate nucleus.
Exceptions to this typical result are included in Table 1.
Note that neocortical subject number 14 had cingulate I
damage in addition to damage characteristic Of
neocortically lesioned animals. Cingulate tissue Spared
in cingulate animals and subject 14 is noted in Table 1.
The amount of cingulate tissue Spared in cingulates varied
from 0.000 cu. mm. to 0.024 cu. mm.

In the case of each Of the fourteen analyses of
behavioral data, a mean score was computed for the three
preOperative sessions. A mean score computed for the
three postoperative sessions was subtracted from the mean
score for preoperative sessions. These difference scores
were analysed according to an unweighted-means two by four
factorial analysis Of variance, the harmonic mean for all
analyses being 4.6602. The mean difference scores are
presented as a function of group membership in Figures 1
through 14.

The results of analyses of behavioral data are
summarized in Table 2. All of the analyses failed to

yield significant E values for lesion group. Analyses of

-|2 1

404

42

 

 

 

 

 

 

 

 

 

 

 

MEAN DIFFERENCE SCORE: FREQUENCY OF MOUNTS
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CINCULATE NEOCCRTICAE 8 H AN N CRMAL
TR E ATMENT

Mean,difference score for preoperative and
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II as a function of treatment group.

43

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r

Figure 2. Mean difference score for preoperative and
postoperative frequency of mounts with intromission

in replications I and II as a function of treatment
group.

44

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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replications I and II as a function of treatment
group.

45

 

 

 

 

 

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46

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Figure 5. Mean difference score for preoperative and
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replications I and II as a function of treatment
group.

47

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.1
200 L.
l.
1 L l l
I l I :
CINCULATE NEOCORTICAL SHAM NORMAL

MEAN DIFFEREME SCORE: LATENCY OF EJACULATION (SEC)

TREATMENT

Figure 6. Mean difference score for preOperative and
postOperative latency of ejaculation in replications
I and II as a function of treatment group.

48

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

430. URI
[Dan
E d
a my
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8
x -40

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II.
E 1 1 J l
O i r 1 l
3 GINOULA‘I’E NEOCORTICAL 3mm NORMAL
g TREATMENT

Figure 7. Mean difference score for preoperative and
postoperative mean interval between mounts in
replications I and II as a function of treatment
group.

49

-500. (II R:

 

 

 

 

 

 

 

 

 

 

; L fill

 

 

 

I 1 l I
I

l I I
Cl NCULATE NEOCORTICAL SHAM NORMAL
T R E A T M E N T

MEAN DIFFERENCE SCORE: KAN NTERVAL em MOUNTS WITH INTRCMISSION (SEC)

Figure 8. Mean difference score for preOperative and
postoperative mean interval between mounts with
intromission in replications I and II as a function
of treatment group.

50

 

 

 

 

 

 

 

 

 

 

 

 

 

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IIIRI
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2
E TREATMENT

Figure 9. Mean difference score for preoperative and
postoperative mean interval between ejaculations in
replications I and II as a function of treatment
group.

51

 

 

 

 

 

 

 

 

 

 

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- [E R1:
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III
’ TREATMENT

Figure 10. Mean difference score for preoperative and
postoperative mean postejaculation interval in
replications I and II as a function of treatment
group.

52

 

 

 

 

 

 

 

 

 

 

 

 

 

 

“-1 DR:
§ ‘ [Dan
E ”m
'2
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g, .,
53-1
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° cINeIILATE NEOOORTIOAL sHAM NORMAL.
3 TREATMENT

Figure 11. Mean difference score for preoperative and
postoperative unadjusted mean number of mounts per
ejaculation in replications I and II as a function
of treatment group.

53

4Q URI
URI:
-Q
_Q 1
_ P_T

O

 

 

 

 

 

 

 

 

 

 

 

 

MEAN DIFFERENCE SCORE: MEAN NMER OF “TS PER EJACULATION (ADJ)
8
N

2d
1
: L I :
CINCULATE NEOCORTICAL SHAM NORMAL
TREATMENT

Figure 12. Mean difference score for preoperative and
postOperative adjusted mean number Of mounts per
ejaculation in replications I and II as a function
of treatment group.

54

 

 

 

 

 

 

 

 

 

 

 

.—
3 4.5. D R I
III R m
3 4.0-.
o
g
F 0.5-
2
E m-
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u 45. ~—
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t I I I L
a cINOULATE NEOOORTIcAI. sIIAM NORMAL
g TREATMENT

Figure 13. Mean difference score for preoperative and
postOperative unadjusted mean number of mounts with
intromission per ejaculation in replications I and
II as a function of treatment group.

55

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Figure 14.

8 URI
"'.5-I
F '1 EUR:
g -mq
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E TREATMENT

Mean difference score for preoperative and

postOperative adjusted mean number of mounts with
intromission per ejaculation in replications I and
II as a function of treatment group.

56

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57

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59

latency to first mount with ejaculation data. frequency
of ejaculation data. and interval between ejaculations
data yielded significant 2 ratios for replication effect.
Each of these replication effects involving a measure of
ejaculation was significant at the .01 level.

Means and standard deviations for the 10 measures of
preoperative and postOperative performance are presented
in Table 3. Table 3 includes means and standard
deviations for the following additional measures of
behavior: mean mounts to ejaculation 1. mean mounts to
ejaculation 2, mean intromissions to ejaculation l. and
mean intromissions to ejaculation 2.

Figure 1 shows that the only subjects failing to
increase the frequency of mounts following surgery were
those of replication I having neocortical lesions. The
greatest increase in frequency of mounts for both
replications occurred in animals subjected to sham
Operation procedures. In marked contrast, Figure 2
suggests that the only subjects failing to decrease the
frequency of mounts with intromission following surgery
were those subjected to sham Operation procedures during
replication 1. The differences in frequency of
ejaculation between subjects in replications I and II.
shown in Figure 3. were large enough to be significant
(£1.31 = 10.320). With respect to subjects showing the
greatest deficits following surgery, the data summarized

in Figures 2 and 3 are similar: i.e.. in both cases the

60

 

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61

 

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62

 

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Table 3 (cont'd.)

Normal

§ham
X

Neocortical

SD

n

SD

n

SD

'2

ingulate

n

SD

Repli- g
X

Measure

cation

 

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Mean mts.

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Mean mts. l-pre
w. intro.

per

ejac.1

63

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64

greatest deficits occurred during replication II in
animals belonging to the neocortical and sham operation
groups.

Figure 0 shows that only slight changes occurred in
latency to first mount for all subjects except those from
replication I having neocortical lesions. the latter
subjects showing a relatively large postOperative
increase in latency to first mount without intromission.
The data summarized in Figure 5 suggest that only those
animals subjected to sham operation procedures during
replication I had shorter latencies to first mount with
intromission after surgery than they did prior to
surgery. The greatest increase in latency to first mount
with intromission occurred in the neocortical lesion
groups. Analysis of the data summarized in Figure 6
indicated that the subjects of replication I differed
significantly (21.31 = 10.866) from those of replication
II with reSpect to the difference in latency to first
ejaculation before and after surgery. The animals in
replication I showed postoperative decreases in latency
to first ejaculation whereas replication II animals
showed postoperative increases in latency to first
ejaculation, the greatest postoperative changes appearing
in the cingulate and neocortical groups.

As shown in Figure 7, the mean interval between
mounts decreased postoperatively for all subjects except

those in the replication I cingulate group and the

65

replication II sham group. the increase for the sham
animals of replication II being slight relative to the
increase shown by the cingulate subjects of replication
I. In contrast to the data presented in Figure 7. the
data summarized in Figure 8 show that the mean interval
between mounts with intromission increased during
postoperative sessions for all subjects except those in
the sham group of replication 1. the greatest postopera-
tive increases occurring in the neocortical groups.
Analysis of the mean interval between ejaculations
difference scores yielded an F ratio significant at the
.01 level (31.31 = 7.978) for replication effects.
Figure 9 clearly suggests that the subjects of
replication I showed a shorter postoperative mean
interval between ejaculations while the replication II
subjects had a longer mean interval between ejaculations
after surgery.

After any ejaculation, regardless of when it
occurred during the 20 minute session, the animal remained
in the observation box until the next mount occurred. The
data presented in Figure 10 suggest that neocortical
animals showed the greatest postoperative increases in
mean postejaculation interval. a similar postoperative
change in the opposite direction being shown by the
replication I sham subjects.

The data presented in Figure 11 represent the

difference scores for unadjusted mean number of mounts

66

per ejaculation. the greatest increases with respect to
this measure of sexual behavior occurring in the normal
groups and the greatest decreases in the neocortical
groups. When the difference score for mean number of
mounts for any subject during any session was based on
only those mounts occurring prior to the last ejaculation
for that session a difference score called the adjusted
mean number of mounts per ejaculation resulted. Figure
12 shows that adjusting the mean difference scores for
mean number of mounts per ejaculation resulted in a
decrease in amount of postoperative change without
altering the basic trends shown in Figure 11.

The apparent differences between replications I and
II with respect to difference scores for unadjusted mean
number of mounts with intromission per ejaculation. shown
in Figure 13. were not statistically significant. Only
the sham animals of replication I and the replication II
neocortical animals failed to conform to the general
trend of postOperative increases in unadjusted mean
number of mounts with intromission per ejaculation for
replication II subjects and decreases in this measure for
replication I subjects. When the score for mean number
of mounts with intromission per ejaculation was adjusted,
as described for adjusted mean number of mounts per
ejaculation, the general trends shown in Figure 13 were
not preserved. As shown in Figure 10. the only subjects

failing to show a decrease in adjusted mean number of

67

mounts with intromission per ejaculation were the
replication II cingulates and the replication I sham
operates.

Due to the infrequency of occurrence of mounts with
intromission not followed by autogenital cleaning, the
autogenital cleaning data were not statistically analyzed.
Only 11 of the males made at least 1. and less than 3,
mounts with intromission without the typical autogenital
cleaning, 7 of these males being observed during
replication I and 4 during replication II. All lesion
groups were represented by the occasional absence of
autogenital cleaning. That is. h of the males were from
the cingulate groups. 2 were from the neocortical groups.
2 were from the replication I sham group, and 3 were from
the normal groups. These males made between an and 115
mounts with intromission during the 6 sessions. 9
responses without autogenital cleaning occurring during
preoperative sessions and 7 responses without autogenital
cleaning occurring during postoperative sessions. ‘No
trends were observable in the mounts with intromission
not followed by autogenital cleaning data.

The eXperimenter was unable to detect signs of motor
deficits in any of the males following surgery.
Observation of qualitative aspects of behavior suggested
that there were only minor postOperative departures from
normal. i.e.. preoperative, sexual behavior. That is.

during the first postoperative session animals 18

68

(cingulate) and 19 (neocortical) tended to fall back and
to the side during autogenital cleaning. Animals 7
(cingulate) and 3 (sham) showed minor departures from the
typical preoperative autogenital cleaning described for
subjects 18 and 19. Notes on behavior, taken by g as
each rat was observed. fail to suggest that the subjects
in either of the lesion groups were unable to carry out
the normal sexual responses in the normal order.

Body weight data collected on alternate days before
surgery and after convalescence from surgery fail to
Support the notion that the animals were sick during
postoperative sessions. No animals weighed less during
the postoperative sessions than they did on the first day
of the eXperiment. Only 2 males in replication I. l
cingulate and 1 normal, showed weight losses between the
last two sessions. Transitory slight weight losses
occurred between the last two sessions in 3 replication
II subjects, 2 cingulates and l neocortical. Because the
subjects were maintained on Wayne Mouse Breeder Blox, it
was occasionally the case that all food particles slipped
through the openings in the bottoms of the cages. Thus,
slight transitory weight losses could be attributed to

the brief absence of food.

DISCUSSION

The results of the present eXperiment appeared to
provide further support for Beach's (1900, 1941) claim
that destruction of less than 20 percent of the cortex
does not result in a disruption of the sexual behavior of
adult male rats. Although the lesion effects reported in
the present eXperiment were not statistically significant.
Figures 2. 3. U, 5, 6, 8, 9, 10, 11. and lb, reflecting
several measures of sexual behavior. showed that
neocortically lesioned rats tended to change more
following surgery than did rats suffering cingulate
lesions. The relatively large behavioral changes obtained
following neocortical lesions might be interpreted as
constituting no evidence contrary to Larsson's (1960)
finding that lateral cortical lesions were more
detrimental with respect to mating behavior in male rats
than were lesions of the median cortex.

The results failed to provide further confirmation
of the work of Stamm (1954), Bunnell and Pinder (1960).
Bunnell. gt_gl. (1966). Stamm (1955), and Slotnick
(1967). More specifically, the results of the reported
eXperiment could not be interpreted as evidence

supporting the hypothesis that the cingulate cortex is an

69

7o

important structure with respect to typical instinctive
behaviors. Since the investigators obtaining evidence
supporting the existence of a relationship between
cingulectomy and behavioral disruptions were not
investigating sexual behavior in the adult male rat.
except for the study of Michal (1965) discussed by Thomas.
Hostetter. and Barker (1968), one might conclude that the
results of the present eXperiment were merely irrelevant
with respect to the instinct studies. Thus the results
appeared to confirm. weakly in the case of Larsson (1964),
the reports of investigations involving sexual behavior
in male rats whereas the studies involving subjects other
than rats and/or non-sexual measures of behavior were not
confirmed. The results reported in this experiment are
also not consistent with the results found by Barker and
Thomas (1965, 1966) in their investigations of acquisition
and retention of alternation behavior following
cingulectomy. Thus, it is concluded that the present
experiment has failed to yield evidence supporting the
hypothesis that mechanisms of temporal-response
integration might be impaired by cingulectomy. It is
conceivable that the small amount of cingulate tissue
spared in most cingulate subjects accounts for the fact
that the present evidence does not confirm the results
reported by Stamm (1950), Bunnell and Pinder (1964).
Bunnell, _e__Ԥ 31- (1966). Stamm (1955). Slotnick (1967).
Michal (1965). and Barker and Thomas (1965, 1966).

71

One might focus upon experimental procedures in
attempting to account for the obtained results. Although
the experimenter intended to use preoperative session one
as a period for selecting responsive males and allowing
the males to gain sexual experience, observations during
the first and second replication I preoperative sessions
resulted in elimination of the extra session. Since all
sessions. including session one, were conducted under
conditions as nearly identical as possible, the observa-
tion that many males showing a high degree of
responsiveness (one or two ejaculations) in the first
session made few or no sexual responses during session
two resulted in no major eXperimental changes.
Replication II was also conducted without the extra
selection and experience session. Because the sessions
lasted only 20 minutes and no rat ejaculated more than
three times per session it does not appear reasonable to
assume that the males had become sexually satiated two or
three days earlier during session one. This conclusion
was based on the results of an investigation of sexual
exhaustion and recovery from exhaustion in the male rat
reported by Beach and Jordan (1956). Beach and Jordan
(1956) found that fully rested males reached the sexual
exhaustion criterion, which consisted of no mounting for
30 successive minutes. after an average of 89.2 minutes
(range of 61.5 to 101.5 minutes). Beach and Jordan

(1956) also found that the mean number of ejaculations to

72

the exhaustion criterion was 6.9 (range of 5 to 10
ejaculations). In terms of the criterion established by
Beach and Jordan (1956), the males in the present
eXperiment had no opportunity to become sexually
exhausted.

One might conclude that the present results do not
constitute sufficient reason for pursuing this line of
research. Although more precise techniques of destroying
brain tissue might lead to more conclusive results, only
minor procedural changes might be in order.‘

Due to severe time and space limitations. only 0 or
5 receptive stimulus females were available for 10 to 12
males on any given day. Although the females were used
no more than three times on a given day and at least 90
minutes intervened between successive pairings, a
preferable procedure would have involved the use of equal
numbers of eXperimental males and receptive stimulus
females.

During pilot observations the eXperimenter was unable
to detect any changes in behavior following the
replacement of the used cobmeal by fresh cobmeal. Since
the females were tested for receptivity soon after the
cobmeal was replaced each day and since all sessions were
preceded by a 10 minute adaptation session, no male rat
was tested in an environment free of olfactory stimuli
from previous rats. A preferable procedure would have

involved the use of fresh cobmeal for each eXperimental

73

rat during each session.

A click occurred each time one of the remote control
panel buttons was depressed. That is, the measurement of
any response was accompanied by a click. Since the
animals were housed in the eXperimental room and in close
proximity to the observation box. this extraneous source
of auditory stimulation was not regarded as an important
variability. However. the use of a silent control panel
would have eliminated the possibility of auditory
contamination.

A further procedural defect involved the Speed at
which the Esterline Angus Recorder was operated. A paper
speed of 3.8 cm. per minute was selected because such a
slow speed eliminated the possibility of running out of
paper while a rat was being observed. This problem could
be eliminated by using a different recording device or by
improving the paper markings. A faster paper Speed would
allow finer time measurements than the four second
minimum used in the present eXperiment.

The 1200 second time limit for latency measurements
during sessions in which no responses occurred was
selected because any given session. with the exception of
longer sessions used to permit measurements of
postejaculatory interval. lasted only 1200 seconds.
Selection of some other latency might have led to
different results.

Although any of these defects might have

7L»

influenced the results, it is unlikely that they would
have obscured large postoperative changes in the sexual

behavior of cingulectomized rats.

APPENDIX A

APPENDIX A

Table 4

Raw data: body weight in grams

 

PreOperative weighings PostOperative weighings

S l 2 2 3

1 443 465 392 435 450
2 325 338 327 357 360
3 365 360 375 384 389
4 353 367 420 421 425
6 405 407 423 437 447
7 428 434 370 418 449
8 354 360 396 407 411
9 331 339 346 356 356
10 332 343 379 392 394
11 375 379 411 416 421
12 353 362 374 377 359
13 348 360 380 377 395
14 359 368 391 397 399
15 366 372 420 413 404
17 352 360 401 408 415
18 334 351 392 393 “03
19 334 348 367 377 380
22 384 394 422 429 438
23 300 312 343 352 351
24 378 378 384 390 383
25 349 365 402 410 414
26 336 342 382 390 394
27 347 363 397 410 419
28 348 363 351 365 345
29 331 340 363 365 372
30 336 371 435 443 455
31 347 368 370 378 364
32 357 361 396 409 409
33 312 326 360 365 369
34 326 335 369 383 402
35 353 364 414 421 431
36 347 349 371 372 382
37 355 358 397 401 408
38 345 358 417 424 429
39 324 325 337 332 336

75

76
Table 4 (cont'd.)

Preoperative weighings Postoperative weighings

 

s 1 2 1 2 3

40 378 384 394 405 409
41 364 374 378 383 393
42 324 334 362 367 379

43 340 341 363 367 386

APPENDIX B

APPENDIX

Table 5

B

Raw data: autogenital cleaning (a) and interresponse times
(irt) in seconds for mounts (m). mounts with intromission
(mi). and mounts with ejaculation (me).

 

 

g and g and g and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
c1ng 2 m 64 m 4 m 48 mia 16 m 72
Rep. 1 mia 4 mia 12 m 32 mia 24 m 120
Pre-l mia 114 mia 44 m 16 mia 28 mia 52
m 48 mia 32 mia 52 m 28 mia 12 m 20
m 8 mia 6O mia 20 m 4 mea 24 mi 8
mia 4 mia 120 mia 28 mia 4 m 424 mia 56
mia 84 mia 118 mia 48 mia 16 m 20 mea 88
mia 84 mia 8 m 48 mia 24 mia 8 mia 188
mia 124 mea 76 mea 4 m 36 mia 24
mia 76 mia 24 mia 28 Pre-2
mia 8O Pre-3 Post-1 mia 28 mia 36 m 24
mea 88 m 4 m 52 mia 4O mia 36 m 4
mia 444 m 8 m 12 m 28 m 36 mia 4
mia 36 mia 4 m 28 mia 28 mia 32 m 8
mia 68 mia 4 mia 4 mia 20 mea 48 m 24
m 8 mia 20 mia 32 mia 8
Pre-2 mia 24 mia 24 mia 48 m 8
m 48 m 16 mia 32 mia 28 cing 6 mia 36
mia 44 mia 12 mia 48 m 28 Rep. 1 mia 24
m 32 mia 20 mia 52 mea 28 Pre-l mia 20
mia 8 mia 16 mia 36 m 268 m 52 mia 24
m 68 m 20 mia 92 mia 216 m 16 mia l6
mia 16 m 36 mea 6O mia 20 m 20 m 20
mia 12 mia 4 m 456 mia 20 mia 8
mia 16 mia 28 mia 56 Post-3 m 12 mia 148
m 56 mia 24 mia 24 m 8 m 24 mia 104
mia 16 mia 20 mia 72 mia 4 mia 12 mia 56
mia 8 mia l6 mia lOO mia 16 m 84 mia 96
mia 20 mia 24 mia 20 mia 4 mia 128
mia 28 m 20 Post-2 mia 36 m 12 mia 16
m 28 mea 8 m 8 mia 32 mia 56 m 48
m 64 m 364 m 64 mia 44 mia 44 mia 32
mia 12 m 20 m 16 mia 28 mia 132 mia 4O
mia 8 m 36 m 68 mia 24 mia 32 mea 4O

77

78

Table 5 (cont'd.)

 

§ and g and g and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
Pre-3 m 24 m 4 mia l6 mia 104 m 36
m 80 mia 24 m 16 mia 28 mia 4O mia 28
m 4 mia 16 m 8 m 32 mia 28 mia 24
m 16 mia 24 m 12 mea 24 mia 8 mia 28
m 32 m 16 m 152 mia 20
m 16 mia 16 m 8 Pre-3 m 20
m 28 mi 12 m 4 cing 7 m 4 m 12
m 4 mia 8 m 4 Rep. 1 m 4 m 8
m 40 mea 12 m 4 Pre-l mia 8 mia 16
m 56 m 184 m 24 mia 48 mia 12 m 40
m 40 m 44 m 52 mia 148 m 144 m 4
m 36 m 32 m 8 mia 140 m 8 mia 4
m 20 m 12 m 12 mia 120 m 96 mea 12
m 24 m 24 m 8 mia 168 mia 12
m 28 mia 72 m 60 m 8 mia 68 Post-2
m 12 m 24 m 128 mia 52 mia 8 m 32
mia 20 m 4 m 4 mia 28 mia 112 m 16
mia l6 mia 4 m 12 mia 80 mia 36 m 4
mia 52 mia 20 m 8 mea 72 m 72 m 4
mia 20 mia 32 m 112 mia 4 m 4
m 24 mia 28 Pre-Z mia 96 m 8
mia 28 mea 24 Post-3 m 28 mia 48 m 20
mia 12 m 4 m 40 mia 12 m 36
mia 36 Post-2 mia 4 m 24 mia 132 m 16
m 24 m 20 m 4 m 52 mea 36 m 40
mia 4 m 4 m 20 m 104 m 36
mia 28 m 12 m 16 m 24 Post-1 m 40‘
mia 36 m 4 m 24 m 28 m 56 m 4
mia 16 m 8 m 16 m 4 m 4 m 16
mia 20 m 32 mia 20 m 4 mia 12 m 52
mia 20 m 12 mia 36 m 44 m 12 mia 4
mia 28 m 12 mia 36 m 8 mia 8 m 4
mea 24 m 4 m 28 m 4 mia 92 m 88
m 248 m 16 m 28 m 72 mia 20 m 16
m 4 mia 12 mi 4 mia 24 m 4
Post-l m 8 mia 28 mia 12 mia 52 m 4
m 48 m 60 mia 12 mia 28 mia 12 mia 4
mia 28 m 32 m 28 mia 20 mia 20 m 32
mia 20 m 48 mea 4 mia 20 mia 16 mia 4
mia 12 m 8 m 192 mia 44 m 16 m 24
m 12 m 12 m 140 mia 128 m 4 mia 8
m 20 m 24 m 28 mia 112 m 8 mia 44
mia 8 m 80 m 20 m 4 m 8 mia 24
mea 24 m 48 m 28 mia 24 mea 4 mia 24
m 168 m 48 mia 4 mia 144 m 284 mia 56
m 76 m 4 mia 24 m 24 m 76 mea 24
m 8 m 16 m 44 mia 4 mia 48

 

 

79

Table 5 (cont'd.)

 

g and g and g and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
Post-3 mia 8 m 24 m 12 m 16 m 92
m 8 m 56 mia 20 mia 44 mia 4 mia 4
mia 4 m 24 mia 4O mia l6 mia 32 m 32
m 8 m 24 m 12 m 12 m 28 mia 4
m 8 m 8 mia 4 mia 12 mia 4 m 52
mia 40 m 56 mia 32 m 20 mia 48 mia 40
m 32 m 144 m 16 m 20 m 20 m 56
m 40 m 16 mea 8 mia 24 m 4 mia 4
mia 16 m 92 m 352 m 16 mia 20 mia 8
mia 48 m 32 m 20 mia 8 mia 32 mia 40
m 20 m 8 mia 12 mia 32 mia l6 mia 48
mia 28 m 60 mia 20 m 24 mia 12 mia 4
mia 32 m 68 mia 28 m 8 mia 24 mia 36
m 24 m 4 mia 40 m 8 mia 12 m 52
mia 8 m 32 mia 20 mea 4 mia 12 mia 28
mia 36 mia 16 mia 20 m 432 mia 16 mia 68
m 20 mia 28 mia 32 m 4 mia 4O mia 24
m 8 mia 24 m 40 m 16 mia 32 mea 32
mea 4 mea 32 m 20 m 32 m 44 m 328
m 372 mia 8 mia 4 mia 20 m 4
m 48 Pre-2 m 20 mia 40 m 36
m 4 m 40 m 8 mia 44 mia 4 Pre-
mia 4 m 20 m 4 mia 20 mia 36 m 20
mia 32 m 40 mia 4 mia 56 mia 28 m 28
mia 44 m 20 m 16 m 28 m 4
m 32 m 4 Post-1 mia 20 mia 8 m 8
mia 24 m 124 m 76 m 24 mia 16 m 24
m 28 m 80 m 32 m 4 mia 24 m 4
mia 32 m 72 m 164 m 8 mia 56 mia 2
m 36 mia 4 m 60 m 4 m 12 mia 5
m 4 m 140 m 444 mea 8 mia 4 m 28
mea 4 m 24 m 140 mea 4 mia 76
mia 12 m 152 Post-3 mia 32
cing 12 m 208 m 16 Pre-2 mia 36
Rep. 1 m 104 m 8 m 100 mia 56
Pre-l m 292 m 96 cing 18 m 8 mia 20
m 156 Rep. 1 m 4 mia 4
m 28 Pre-3 Post-2 Pre-l m 4 m 40
m 44 m 4 m 88 m 124 m 4 mia 4
m 42 mia 8 mia 4 m 4 mia 28 m 12
m 24 mia 24 mia 20 mia 20 m 4 m 88
mia 84 mia 20 m 20 mia 24 mia 8 mia 4
m 8 m 12 m 12 m 16 m 24 m 24
m 4 mia 8 mia 12 mia 20 m 4 mia 4
m 60 mia 20 mia 24 m 52 m 16 m 16
m 4 mia 28 mia 24 mia 24 m 28 mea 8

80

 

 

Table 5 (cont'd.)

g and § and g and g and g and § and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 328 Post-2 Pre-l m 12 m 432 m 168
m 12 m 20 m 52 mia 4 m 4 m 12
m 116 m 28 mia 36 m 16 mia 4 mia 12
m 12 m 44 m 12 mia 32 mia 12 mia 168
m 4 m 656 mia 356 mia 24 mia 20 mia 64
mia 4 m 32 mia 124 m 24 mia 4O mia 120
m 24 m 148 mia 16 mia 52 mea l6 mea 32
m 8 m 44 mia 112 mia 8 m 308
mia 4 m 12 mia 224 mia 4 Post-2
mia 32 m 4 mea 28 m 20 m 40 Pre-2

m 32 mia 64 m 4 m 4 m 140
Post-l m 84 mia 8 mea 20 m 16 m 4
m 32 m 8 m 16 m 384 m 32 m 96
m 4 mia 12 m 20 m 32
m 4 Post-3 m 36 Post-3 m 32
mia 4 m 48 Pre-Z m 4 mia 72 m 80
mia 24 mia 12 m 64 mia 4 mia 48 m 100
mia 32 mia 8 m 164 m 40 mia 12 m 172
mia 48 m 16 m 788 m 8 mia 32 mia 4
mia 28 mia 16 m 136 m 4 mia 48 m 24
mia 24 mia 20 m 4 mia 8 mia 44 m 244
mia 4O mia 16 m 28 mia 24
mia 24 mia 20 Pre-l mia 16 m 32 Pre-3
mia 36 mia 20 m 8 m 16 m 24 m 8
mia 16 mia 12 mia 8 m 12 m 12 m 96
m 28 mia 12 mia 8 mia 8 mia 8
mea 4 mia 20 m 20 Post-l m 40 mia 24
m 152 mea 12 m 4 m 40 mia 12 m 24
m 32 m 332 mia 20 mia l6 mea 32 mia 12
m 32 m 28 m 16 mia 20 mia 468 mia 28
m 52 mia 11 mia 28 mia 12 m 20 mia 20
m 32 mia 24 m 20 mia 12 m 40 m 12
m 4 mia 28 mia l6 mia 32 mia 8 mia 24
m 4 mia 20 mia 16 mia 16 mia 20 m 20
mia 4 mia 24 m 8 m 16 mia 36 m 24
mia 52 mia 20 mia 12 mia 12 mia l6 mia 16
mia 48 m 16 m 12 m 32 m 48 mia 36
mia 36 mia 8 mia 12 mea 8 mia 36 mia 16
mia 44 m 24 m 44 mia 344 mia 52
mia 32 mea 4 mia 4 m 12 m 276
mia 40 m 280 mia 12 mia 8 neo 14 m 20
m 24 m 60 mia 16 m 12 Rep. 1 m 20
m 16 m 60 mia 12 Pre-l m 24
mea 12 mia 4 mia 16 m 188 mia 8
m 180 neo 9 mia 20 m 16 m 108 mia 24
m 72 Rep. 1 mia l6 mea 12 mia 4 m 36

81

Table 5 (cont'd.)

 

§ and § and g and § and g and
beh. irt beh. irt beh. irt beh. irt beh. irt
m 28 mia 12 mia 44 mia 6O Pre-3
m 16 mia 88 mia 40 m 48
m 4 mia 28 mia 36 m 24
neo 19 m 48 m 284 m 24
Rep. 1 mia 8 Post-2 mia 68 m 8
Pre-l m 40 m 16 mia 28 m 40
m 84 mia 4 m 12 mia 36 m 48
m 40 mia 36 m 24 mia 36 m 100
mia 4 m 88 m 40 mia 32 m 148
m 64 mia 4 m 44 mia 32 m 56
m 4 mia 52 m 12 mia 52 mia 76
mia 4 mia 60 m 52 mia 28 m 24
mia 108 mia 60 m 60 mia 24 mia 12
mia 6O mea 24 m 24 mia 36 mia 12
mia 128 m 328 m 4 mia 36 mia 4O
mia 116 m 228 m 76 mia 24 m 16
mia 96 mia 4 m 40 mia 32 mia 12
mia 104 m 48 mia 4 m 24 m 20
mia 144 mia 12 m 48 mia 24 m 4
m 8 mia 108 m 4 mia 20 m 24
mea 96 Post-l m 12 mia 24 m 8
m 8 mia 4 mia 8
Pre-2 m 12 m 28 mia 56
m 8 mia 8 m 4 sham l mia 28
mia 16 m 108 mia 56 Rep. 1 m 40
m mia 36 m 8 mia 24 Pre-l mea 4
m mia 44 m 32 mia 20 m 108 m 216
m mia 24 mia 4 mia 28 m 28 m 24
m mia 8 mia 16 mia 24 m 44 Post-1
m mia 28 mia 32 mia l6 mia 84 m 20
m mia 52 m 56 mia 28 mia 36 m 20
m mia 72 mia 4 mea 24 mia 92 m 16
m mia 36 mia 68 m 264 mea 48 m 44
m mia 44 mia 44 m 4 mia 456 m 40
m mia 84 mia 40 m 56 mia 36 m 40
m m 32 m 40 m 4 mia 84 m 24
mia 68 mia 68 m 92 mia 152 m 4
P mea 64 mia 36 mia 4 mia 76 mia 8
m m 468 mea 56 mia 24 mea 20 mia 28
m mia 8 m 188 mia 36
m mia 44 m 140 Post-3 Pre-2 m 16
m m 4 mia 4 m 68 mia 20
m Pre-3 m 4 mia 28 m 184 m 16
m m 4 m 32 mia 44 m 700 mia 16
m mia 8 mia 4 mia 48 m 144 m 32

82

Table 5 (cont'd.)

 

§ and g and g and S and S and § and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 8 mia 8 m 16 mia 28 Post-2 mia 20
m 4 m 16 m 20 mia 24 m 20 mia 32
mia 20 mia 8 m 12 mia 32 m 8 mia 36
mia 48 mia 24 m 16 mia 12 mia 4 mia 40
m 16 m 12 m 20 mia 20 mia 20 m 36
m 12 mia 12 m 28 mia 28 mia 20 m 4
mia 20 m 12 m 20 mia 12 mia 28 mea 4
m 12 mia 20 mia 20 mia 32 m 20 mia 528
mia 4O mia 20 m 16 mia 12 mia 8 m 12
mea 20 m 12 m 8 mia 20 m 16 m 12
m 240 m 20 m 16 m 20 m 12 m 8
m 168 m 20 m 16 mia 24 mia 4 m 20
mia 4 mia 20 m 32 mia 48 mia 16 mia 16
m 20 m 36 mia 12 mia 16 m 28 m 24
mia 36 m 12 m 4 mea 44 mia 16 mia 24
mia 24 m 24 mea 4 m 180 m 16 m 20
m 32 m 16 m 56 m 12 mia 12 mea 4
m 28 m 20 mia 92 m 12
mia 20 mia 28 m 28 m 40
m 28 m 16 sham 3 mia 32 m 20 sham 10
m 4 mia 24 Rep. 1 mia 16 m 4 Rep. 1
m 4 m 20 Pre-l mia 24 mia 8 Pre-
m 24 m 28 m 56 mia 16 m 16 m 200
m 28 m 12 m 72 m 24 m 116 mia 40
m 28 m 24 mia 52 mea 4 m 20 m 16
m 12 mia 8 mia 88 m 148 m 208 m 48
m 56 m 16 mia 44 m 12 m 108 m 16
m 68 m 8 mea 76 m 88 m 52 m 28
m 96 mia 16 mia 432 m 96 mia 24 m 28
m 28 m 20 mia 68 mia 20 m 24
m 64 mia 20 mia 92 Post—l mia 12 mia 80
m 124 m 20 mia 52 mia 24 m 12 m 12
m 28 mia 28 mea 76 mia 8 mia 20 mia 44
m 48 m 28 mia 28 m 20 mia 28
m 12 mia 12 Pre-2 mia 56 m 8 m 28
m 212 m 20 m 56 m 40 m 16 m 20
m 20 m 20 m 28 m 16 mia 28 m 108
m 84 mia 12 m 12 mia 44 m 24 mea 32
m 44 m 16 m 288 m 304 mea 8 m 228
m 36 m 20 m 536 mia 184
m 44 m 20 mia 44 Post-3 Pre-2
m 120 m 4 Pre-3 mia 32 m 4 m 12
m 24 m 4 m 52 mia 8 m 80
Post-3 m 24 m 4 m 8 mia 12 m 92
mia 4 m 8 mia 12 mea l6 mia 20 m 40
m 8 m 20 mia 28 m 16 m 124

83

Table 5 (cont'd.)

 

g and g and g and § and § and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 460 m 132 mia 28 mia 24 m 12 mia 28
m 4 m 4 m 72 m 24 mia 8 mia 48
m 36 m 132 mia 4 mia 20 m 16 mia 24
mia 4 m 44 m 12 mia 16 m 12 m 20
mia 68 m 24 m 16 m 24 m 4 mia 8
mia 36 mia 16 mea 8 mea 12 mia 8 m 12
mia 16 m 16 m 240 m 344 m 24 mia 28
m 36 m 36 m 32 mia 12 mia 4 mia 16
mia 28 mia 4 m 20 mia 12 m 20 mia 16
mia 28 m 20 m 64 mia 24 mia 32
mia 32 m 32 m 20 Post-3 m 16 mia 36
mia 36 m 20 mia 72 mia 4 mia 4 mia 24
mia 64 mia 4 m 8 m 8 m 32 mea 52
mia 48 mia 12 m 8 mea 4 m 372
Pre-3 m 20 mia 4 m 12 mia 8
m 8 m 12 m 24 mia 12 mia 52
m 8 m 44 m 56 mia 8 sham 22 mia 16
m 4 m 8 mia 4 mia 12 Rep. 1 mia 24
mia 12 mia 4 m 52 m 24 Pre-l mia 16
m 12 m 20 mia 8 m 4 m 36 mia 40
m 12 m 24 m 8 mia 4 m 24 mia 20
m 4 mia 32 mia 4 m 20 m 64 mia 8
mia 12 m 20 m 96 m 4 m 4 mia 6O
mia 12 mia 8 m 24 mia 6O mea 32
m 20 mia l6 Post-2 m 4 mia 28
m 20 m 16 m 4 m 12 mia 20 Pre-3
mia 28 m 24 mia 12 m 4 mia 88 m 4
mia 16 mia 8 mia 12 mia 28 m 4
m 16 Post-1 mia 12 m 12 mia 20 mia 12
m 8 m 12 mia 16 m 12 mia 84 m 8
m 12 m 24 mia 20 m 4 mia 4 mia 4
m 8 mia 12 mia 60 m 12 mia 4 m 20
mia 4 mia 8 mia 16 m 4 m 72 mia 16
m 28 mia 12 mia 16 m 4 mea 20 m 24
mia 4 m 20 m 20 mia 8 mia 448 m 20
m 20 mia 20 mia 20 m 16 mia 52 mia l6
mia 4 m 20 mia 20 mia l6 mia 100 mia 20
mia 20 mia 36 mia 16 m 28 mia 24
m 20 m 8 mia 20 mea 4 Pre-2 mia 24
m 8 mia 8 m 24 m 256 m 8 mia 16
mia 28 m 20 mia 20 m 28 m 8 m 20
m 44 m 28 mia 32 m 116 m 4 m 16
m 12 m 16 m 20 m 4 mia 4 mea 4
mia 12 m 32 mea 52 m 4 mia 16 m 252
mia 4 m 16 mia 308 mia 16 m 28 mia 4
mea 28 mia 4 mia 12 m 12 mia 4O mia 28

84

Table 5 (cont'd.)

 

g and § and § and § and § and
beh. irt beh. irt beh. irt beh. irt beh. irt
m 16 mia 36 m 8 m 28 nor 8
m 4 m 12 m 32 Rep. 1
mea 4 mea 24 m 4 Post-l Pre-l
m 192 m 236 m 4 m 8 m 16
m 76 m 4 m 16 m 24 m 8
m 12 m 32 m 172 m 12
Post-2 m 76 m 4 mia 8
m 12 m 80 m 12 Post-2 mia 28
m 28 m 4 m 40 m 56 m 20
m 12 mia 4 m 40 m 32 mia 32
m 4 mia 36 m 32 m 68 mia 160
m 8 m 20 m 8 m 76 mia 16
m 16 mia 24 m 28 m 48 mia 32
m 16 m 28 m 100 m 20 mia 316
mia 8 mia 20 m 228 m 28 mia 24
m 20 mia 28 m 16 m 4 m 28
mia 4 m 24 m 36 m 52 mia 68
m 28 m 4 mia 4 m 24 mea 92
mia 4 mea 4 m 8 m 160 mia 104
mia 32 m 212 m 144 m 404 mia 160
m 16 m 92 m 320
m 4 Post-3 Pre-2
mia 8 Pre-3 m 4 m 4
mia 36 nor 4 m 4 m 4 m 12
m 20 Rep.1 m 4 m 8 m 40
m 4 Pre-l m 4 m 20 m 8
mia 4 m 148 m 4 m 44 m 8
m 296 mia 8 m 4 m 52 m 32
m 24 mia 112 m 24 m 16 m 24
m 48 mia 72 m 8 m 12 m 44
mia 4 mia 56 m 4 m 28 m 12
mia 44 mia 32 m 28 m 56 m 12
m 16 mia 40 m 152 m 16 m 4
mia 16 mia 28 m 116 mia 4 m 24
mea mia 28 mia 72 m 48 m 20 m 4
m mia 20 mia 56 m 344 m 4 m 104
m m 28 mia 24 m 4 m 40 m 76
m mea 8 mia 20 m 20 m 4 m 144
m mia 28 m 4 m 84 m 16
m Post-3 mia 100 m 28 m 16 m 32
m mia 8 mia 48 m 20 m 108 m 8
mia mia 20 mia 32 m 36 m 48 m 72
m mia 16 mea 36 m 88 m 288 m 84
mia m 20 m 24 m 4 m 12
m mia 4 Pre-2 m 44 m 112 m 44
mia mia 24 m 16 m 96 m 100

85

Table 5 (cont'd.)

 

g and S and § and S and S and S and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 32 m 16 m 12 m 20 Pre-2 mia 32
m 56 m 28 mia 12 m 12 m 24 mia 32
mia 4 m 36 m 16 mia 12 m 12 mia 104
mia 44 m 8 m 16 mia 12 mia 4 m 40
Pre-3 m 4 mia 8 m 24 m 4 m 8
m 4 m 12 mia 12 mia 24 m 16 m 72
m 4 m 4 mia 32 mia 12 mia 16 m 4
mia 4 m 16 mia 24 m 32 mia 88 mea 4
m 80 m 28 m 32 m 8 m 4
mia 52 m 4 m 28 m 4 mia 4 Post-1
mia 72 m 36 m 36 mia 8 mia 32 m 84
mia 80 m 48 mia 12 m 36 mia 88 m 20
mia 40 m 8 m 56 mea 4 mia 140 m 56
m 48 m 4 mia 4 m 324 mia 112 m 4
mia 28 m 4 mia 16 m 44 mia 68 m 72
mia 32 m 16 mia 48 m 76 mia 52 m 4
mia 48 m 4 m 40 m 4 mia 132 m 84
mia 44 m 4 mia 8 m 4 m 8 m 32
m 20 m 4 m 60 m 44 mia 4 m 4
mia 12 m 12 mia 4 m 4 mia 172 m 4
mia 56 mia 4 mia 16 mia 4 m 108
m 28 m 28 m 88 mia 28 Pre-3 m 204
mia 12 mia 8 mia 4 mia 48 m 16 m 156
mia 44 m 24 m 56 mia 28 m 32 m 24
mia 68 mia 4 mia 4 mia 60 m 24 m 168
mia 12 m 36 mea 12 m 36 m 20 m 4
m 16 mia 16 m 172 m 20 m 76
mia 48 m 24 m 72 mia 4 mia 4 Post-2
m 28 mia 8 m 236 mia 4 m 20
mia 8 mia 20 m 4 m 8 m 32
mia 48 mia 36 mia 4 nor 11 m 16 m 4
m 44 m 36 m 32 Rep. 1 mia 4 mia 4
mea 60 mia 4 mia 12 Pre-l mia 36 mia 16
m 16 m 96 m 24 m 8
Post-1 m 24 Post-3 m 56 m 8 mia 4
m 4 m 8 m 4 mia 16 mia 4 m 20
m 4 mia 4 m 4 mia 8 m 56 m 4
m 4 m 56 m 20 m 64 mia 40 mia 36
m 12 mia 8 mia 8 m 76 mia 72 mia 12
m 28 mia 12 m 40 m 52 mia 20 mia 20
m 40 m 36 m 60 mia 64 mia 60 mia 32
m 8 mea 4 mia 4 mia 96 mia 36 mia 24
m 24 m 128 mia 28 mia 16 mia 32 mia 40
m 20 mia 20 mea 16 mia 20 mia 24
m 48 Post-2 m 48 mia 300 mia 48 mia 28
m 24 mia 4 mia 20 mia 132 m 20 mia 4O

86

Table 5 (cont'd.)

 

S and § and g and § and g and § and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 16 m 20 mia 36 m 36 m 8 mia 4
m 32 mia 4 mia 28 m 40 m 8 m 24
mia 36 m 20 m 56 mia 24 mia 8 m 12
mea 32 mia 4 mia 4 mia 48 m 8
m 320 mia 36 mia 4O mia 24 m 8 Post-3
mia 8 m 20 m 60 m 28 m 12 m 4
m 24 m 4 m 12 mia 8 m 8 m 4
mia 12 m 4 m 12 m 24 m 12 m 4
mia 32 m 4 mea 12 m 20 mia 16 m 4
mia 44 m 4 m 440 mia 4O mia 4O mia 4
mia 36 mea 4 mia 4 m 48 mia 24 m 28
mia 4O mia 52 m 4 mia 32 mia 20
m 28 mia 44 m 4 mia 20 mia 32
m 4 nor 13 mia 48 mea 4 mia 24 m 36
mea 4 Rep. 1 m 60 mia 36 mia 48
Pre-l m 4 Post-l m 20 m 4
Post-3 m 20 m 8 mia 16 mia 12 mia 4
m 4 m 36 mia 20 mia 16 m 24 mia 52
m 4 m 24 mia 20 mia 24 m 4 m 28
mia 4 m 8 m 36 m 32 mia 8
m 16 m 40 Pre-3 mia 4 m 20 m 24
mia 20 m 80 m 4 m 12 m 16 m 20
mia 12 m 104 m 12 m 4 m 12 m 48
mia 20 mia 4 mia 4 m 32 m 4 m 4
mia 28 mia 88 m 24 mia 4 m 4 mia 36
mia 28 mia 4 mia 4 mia 12 m 4 m 32
mia 20 mia 28 m 24 m 20 m 4 m 16
m 20 mia 4O mia 32 m 12 mea 4 mia 40
m 20 mia 72 mia 44 mia 16 m 384 m 28
mia 4 mia 12 m 24 m 20 m 28 m 4
mia 20 mia 68 mia 24 mia 8 mia 4 m 4
mia 24 mia 72 m 24 m 44 m 40 m 28
m 28 mia 56 m 24 mia 8 m 20 m 36
mia 36 m 92 m 4 m 28 mia 4 mia 4
m 12 mia 4 mia 16 mea 4 mia 28 mea 4O
mia 4 mia 48 m 68 m 396 mia 36 m 396
m 28 mea 40 m 8 mia 4 m 56 m 4
mia 16 m 20 m 12 mia 4 m 16
m 320 Pre-2 m 4 mia 4 m 24 m 4
m 12 m 36 m 8 mia 20 m 4 m 24
m 36 m 32 m 4 m 16 m 4 mia 20
mia 4 mia 4 mea 8 mia 24 m 4 mia 36
m 16 m 28 m 356 mia 36 m 4 m 40
m 8 mia 40 m 52 mea 24 m 4
mia 4 m 36 mia 4 m 4
mia 36 mia 20 mia 36 Post-2 m 4 nor 15

87

Table 5 (cont'd.)

 

§ and § and § and § and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
Rep. 1 m 16 m 20 mia 4 m 28 mia 20
Pre-l mia 4 mia 8 mia 16 mia 8 mia 48
m 432 m 28 mia 28 mia 8 mia 76
mia 76 m 12 mia 12 Post-3 mia 8 mia 28
mia 64 mia 8 mia 52 m 12 mia 16 mia 20
mia 132 mia 24 m 12 m 4 mia 24 mia 24
mia 48 m 24 m 4 mia 4 mia 24 mia 4
mia 76 mia 24 mia 4 m 16 m 12 mia 36
mea 160 mia 16 m 20 m 4 mia 28 mia 48
mia 196 mia 56 mia 8 mia 8 mia 44 mia 56
mia 28 m 16 mia 8 mia 28 mea 24
Pre-Z mea 44 mea 4 m 20 mia 88 m 236
m 16 mia 492 mia 8 mia 16 m 8
m 4 m 4 Post-2 mia 8 mia 4 m 48
m 4 mia 4 m 28 mia 28 mia 4 m 48
m 20 mia 40 m 4 m 32 mia 28 mia 12
m 8 mia 52 m 48 mia 8 mia 36 mia 12
m 20 m 12 m 4 mia 44 mia 20 mia 36
m 48 mia 68 m 4 mia 16 mia 8 mia 24
mia 4 mia 20 m 4 mia 24 mea 20 mia 8
m 16 mia 28 m 4 mea 32 m 280 mia 28
m 32 mia 32 mia 4 m 412 m 4 mia 12
m 68 mea 40 m 40 m 64 mia 28 mia 12
mia 36 m 4 m 4 mia 28 m 24
m 32 Post-l m 80 mia 4 mia 12
m 4 m 8 m 16 mia 20 mia 8 Pre-3
mia 4 mia 4 m 160 mia 36 mia 32 m 4
m 8 m 52 m 52 mia 24 mia 16 m 4
mia 4 m 16 m 4 mia 24 m 16 m 4
m 44 mia 4 mia 4 mia 20 mia 4 m 4
m 228 mia 24 mia 16 m 28 m 12
mia 4 m 32 m 24 m 16 Pre-2 mia 8
m 104 mia 4 mia 4 mia 36 m 24 mia 12
m 16 mia 12 mia 20 mia 16 m 8 m 28
mia 4 mia 20 mia 16 mia 40 m 4 m 24
m 164 mia 28 m 16 m 48 m 4 m 4
m 120 mia 32 mia 4 mea 20 m 16 mia 8
mia 52 mia 20 m 4 mia 56
Pre-3 m 20 m 16 m 20 mia 28
m 8 mia 12 m 12 nor 17 m 20 mia 56
mia 4 mia 52 mia 16 Rep. 1 m 8 m 60
m 12 mia 16 mia 36 Pre-l mia 12 mia 4
m 24 mia 16 m 16 mia 232 mia 68 mia 8
mia 8 mea 16 mea 4 m 28 m 72 mia 12
m 24 mia 496 m 488 m 44 mia 4 mia 20
mia 12 mia 8 m 4 m 20 m 32 mia 4

88

Table 5 (cont'd.)

 

§ and g and g and § and § and S and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mea 4 m 44 mia 4 mia 6 mia 32 m 8
mia 508 mia 24 m 24 mia 4 mia 20 mia 8
mia 16 mia 16 m 12 mia 112 mia 20 m 36
m 44 mea 24 mia 16 mia 40 m 16 mea 4
mia 8 m 68 mia 44 mia 12 m 352
mia 44 Post-2 mia 4 mia 88 m 12 mia 4
mia 12 m 4 mia 16 mea 80 m 24 mia 20
m 12 m 4 m 48 m 272 mia 12 m 20
mia 36 m 8 mia 4 m 4 mia 12 m 8
mia 20 m 8 mia 36 m 28 mia 12 m 16
mia 16 m 24 mia 20 mia 60 m 36 mia 12
m 20 m 8 m 20 mia 8 m 4 mia 20
mia 44 m 4 mea 4 mia 24 mea 4 m 20
mia 4 m 4 m 20 m 412 m 12
m 16 m 4 Post-3 m 12 mia 4 m 16
mia 4 m 16 m 8 mia 4 mia 16 m 4
m 4 mia 4 m 16 m 4
Post-1 m 4 m 12 Pre-2 mia 8 m 48
m 4 m 20 mia 12 mia 4 mia 44 m 8
mia 4 m 8 mia 20 mia 12 m 12 mea 8
mia 8 m 16 m 68 mia 12 m 12 m 320
m 12 m 20 mia 28 m 16 m 16
m 12 m 36 mia 20 m 12 mia 8 Post-2
m 4 m 8 mia 24 mia 12 mia 36 m 28
m 44 m 16 mia 24 m 12 m 24 m 60
mia 4 m 4 mea 24 mia 12 mia 8 m 52
m 8 m 12 m 120 mia 20 m 24 m 200
mia 32 m 116 m 60 mia 24 mia 16 m 44
m 24 m 36 m 96 mia 48 m 16 m 268
mia 4 m 20 m 120 mea 32 m 8 m 132
mia 24 m 24 m 24 m 272 mea 8 m 4
mia 44 m 4 mia 48 mia 4
mia 40 m 20 mia 44 m 16 Post-1 Post-3
mia 40 m 44 mia 44 mia 16 mia 8 mia 4
m 16 m 4 mia 28 mia 20 m 20 m 8
m 60 m 20 mea 32 mia 6O mia 24 mia 8
mia 12 m 16 m 256 mia 16 m 16 mia 16
mia 20 m 28 m 24 mia 16 m 36
mia 24 m 52 mea 44 m 28 mia 36
mea 20 m 24 cing 24 m 12 mia 40
m 452 m 44 Rep. 2 Pre-3 mia 4 mia 84
mia 8 m 4 Pre-l mia 8 m 20 mea 64
mia 20 m 60 m 8 mia 28 mia 8 m 280
mia 16 m 36 m 4 m 24 mia 28 mia 8
mia 24 m 8 mia 24 mia 4 m 16 mia 16
mia 24 m 4 mia 64 mia 20 m 4 mia 28

89

Table 5 (cont'd.)

 

 

g and g and g and g and g and § and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 20 mia 28 m 8 mia 8 m 44 mia 20
mia 16 mia 16 mia 8 mia 4 m 24
m 24 mia 24 mia 16 m 12 m 24
mea 8 mia 24 mia 24 m 4 cing 29 mia 16
m 276 m 20 mia 20 m 4 Rep. 2 mea 24
m 128 mea 20 mia 20 m 8 Pre-l m 180
m 20 mia 356 m 20 mia 12 m 16 m 384
m 4 mia 8 m 8 m 16 mia 20 mia 4
mia 24 m 16 mia 16 mia 16 mia 12 mia 20
mia 12 m 16 m 20 mia 20 mia 24 mia 24
mia 24 mia 16 m 16 mia 16 m 16 m 24
m 20 m 8 mia 16 mia 12 mia 20
mia 8 mia 12 mia 12 mia 20 mia 20
cing 28 m 16 mia 24 mia 8 mia 16 m 28
Rep. mea 4 mia 20 mia 16 mia 20 mia 20
Pre-l m 116 m 16 mia 16 mia 16 m 64
m 40 m 12 mia 16 m 12 mia 20 mia 16
mia 4 mia 16 mia 24 mia 4 m 16
m 28 mia 16 mia 24 mea 16 mea 20 Pre-3
m 28 mia 8 m 24 m 124 mia 456 m 16
mia 12 mia 20 m 4 m 136 mia 20 mia 4
mia 36 m 28 m 4 mia 16 mia 4
m 24 mea 8 mea 4 Post-3 mia 24 mia 20
mia 4 mia 356 m 4 mia 16 mia 24
mia 36 Pre-3 m 16 m 8 m 16 mia 24
mia 28 m 32 mia 12 mia 4 mia 20 m 24
m 36 m 132 mia 12 mia 20 m 28 mia 40
mia 4 m 76 m 12 mia 12 mia 20 mia 16
m 32 m 20 m 12 m 16 mea 20 m 20
mea 32 m 88 m 4 mia 8 m 312 mia 4
m 316 m 352 m 4 m 12 m 48
mia 36 m 312 mea 20 mia 12 Pre-2 mia 4
mia 16 m 200 m 12 m 8 mia 16
m 16 m 4 Post-2 mea 4 mia 4 mia 36
mia 8 m 12 m 4 m 240 mia 16 mia 12
mea 20 m 16 m 100 mia 16 mea 16
m 372 Post-1 m 12 m 60 mia 36 m 580
mia 56 m 8 m 36 mia 4 mia 20 mia 4
mia 8 m 104 m 20 mia 16 mia 20
Pre-2 mia 12 m 36 mia 8 mia 16 m 20
m 8 mia 12 m 32 mia 44 mia 20 mia 4
mia 4 m 8 m 84 m 24 mia 4O mia 16
m 8 mia 4 m 64 mea 8 mia 16 mia 20
mia 16 mia 16 m 72 m 320 mia 28 mia 16
mia 16 mia 12 m 44 m 84 mia 20 mia 24
mia 12 m 16 m 96 m 112 mia 24 mia 16

9O

 

 

Table 5 (cont'd.)

g and § and g and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 12 m 56 mia 48 mia 8 mia 64 m 4
mia 28 m 16 mia 168 mia 60 m 40 mia 4
m 20 m 52 mia 148 mia 44 mia 4O mia 84
mia 8 m 232 mia 188 mia 44 mia 20 m 8
mea 20 m 12 mia 216 mia 96 mia 32 mia 56

m 472 mia 64 mea 24 mia 64 m 8
Post-1 m 104 mia 144 mia ' 448 mia 24 mia 4
m 4 m 80 m 48 m 16 mia 32 mia 28
mia 20 mea 4 mea 32 mia 28
mia 16 Post-3 Post-1 m 372
mia 20 m 4 Pre-Z m 8 mia 8 Pre-2
m 16 mia 4 m 4 m 4 mia 4 m 4
mia 4 mia 28 m 4 mia 32 mia 68 mia 4
mia 16 mia 48 mia 8 mia 6O mia 100 mia 20
m 36 mia 20 m 48 mia 6O mia 12
mia 12 m 40 m 16 mia 40 m 36
mia 16 mia 4 m 60 mia 6O cing 39 mia 20
mia 28 mia 44 mia 12 mia 68 Rep. 2 mia 20
m 28 mia 36 mia 72 mia 64 Pre-l mia 24
mia 36 mia 28 mia 208 m 28 mia 44 mia 28
mia 24 m 48 mia 60 m 8 m 16 mia 60
mia 16 m 60 mia 112 mia 4 mia 32 mia 36
mia 36 mea 4 mia 4O mia 44 mia 20 mia 32
m 16 m 188 mia 32 mia 48 mia 20 mia 36
mia 28 m 40 mia 120 m 44 m 24 m 20
mia 20 m 16 mia 16 mia 24 mia 4 mia 24
mia 28 m 44 mia 6O mea 68 mia 20 mia 36
mia 24 m 36 mia 96 m 376 m 60 mia 24
mia 48 m 112 mia 4O mia 2O mia 4 mia 28
m 12 m 12 mia 48 mia 32 mia 48 m 20
mia 40 m 20 mia 24 mia 84 m 12 mia 36
mia 16 mia 28 mia 92 mia 4 mia 20
mia 28 mia 84 m 24 Post-2 mia 28 mea 40
m 16 mia 48 m 8 mia 16 m 408
mia 16 mia 44 Pre-3 m 20 mia 36 mia 12
mia 16 m 72 mia 4 m 24 mia 24
mea 12 mia 12 mia 128 Post-3 m 4 mia 32
m 284 mea 40 m 32 m 4 mia 68 mia 20
m 160 m 44 mia 4 m 12 mia 20
mia 4 mia 8 m 36 mia 4 mia 48
mia 28 cing 34 mia 6O mia 16 mia 32 mea 20
mia 16 Rep. 2 m 4 mia 40 mi 12
mia 16 Pre-l mia 56 mia 52 mea 4 Pre-3

m 8 m 24 mia 12 m 372 m 4
Post-2 m 4 m 72 mia 36 m 4 m 4
m 4 m 4 m 20 mia 32 m 32 mia 12

91

Table 5 (cont‘d.)

 

§ and g and § and S and § and S and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 12 mia 32 mia 20 m 4 mia 1068 Pre-2
mia 20 mea 28 m 24 mia 16 m 8
mia 16 mia 232 mia 24 m 12 Post-2 m 4
mia 12 mi 4 mia 40 mia 12 m 20 mia 12
mia 24 m 28 mia 24 m 24 mia 12
mia 20 Post-2 mea 28 mia 24 m 120 m 16
mia 20 mia 12 m 464 mia 20 m 256 mia 16
mia 32 mia 8 m 36 mia 28 mia 36
mia 20 mia 16 mia 4 mia 24 Post-3 mia 28
m 36 m 12 mia 20 mia 24 m 692 m 20
m 4 mia 16 mia 20 m 16 m 360 m 4
mia 4 m 24 mia 24 mia 24 mia 4
m 28 mia 16 mia 24 mia 16 m 40
mea 4 mia 24 mia 36 m 16 neo 31 mia 4
m 412 mia 16 mia 24 mia 20 Rep. 2 m 24
mia 4 mia 52 mea 48 mia 24 Pre-l m 24
mia 48 mia 32 m 24 m 12 mia 4
m 32 mia 16 mia 28 m 8 mia 24
mia 24 mia 20 neo 27 mia 32 mia 12 m 40
mia 56 mia 36 Rep. 2 m 24 mia 16 mia 36
m 28 mia 20 Pre-l mea 4 mia 24 mia 20
mia 4 mia 28 m 36 mia 436 mia 32 m 28
mea 44 mea 44 mia 4 m 28 mia 44 m 8
m 164 mia 6O mia 12 mia 24 mia 4
Post-1 m 52 m 20 mia 4O mia 32 m 16
m 8 m 48 m 52 m 24 mia 24 mea 4
mia 12 m 88 mia 4 m 12 mia 24 m 416
mia 24 m 20 m 28 mia 52 mia 24 m 4
mia 16 mia 48 mia 36 mia 40 mia 32 m 12
mia 20 mia 24 mia 48 m 24 mea 40 m 28
mia 32 mia 20 m 36 mea 24 m 308 mia 44
mia 28 mia 20 mia 16 m 4 mia 32
mia 20 mia 20 mia 76 Pre-3 m 4 mia 32
mia 16 mia 32 mia 20 m 24 mia 24 m 100
mia 32 m 24 m 52 m 12 mia 44 mia 40
m 20 mia 16 mia 4 m 20 mia 12
mia 24 mia 28 mia 56 m 4 mia 32 Pre-3
mia 32 mia 24 m 20 m 32 mia 32 mia 24
m 32 mea 44 m 8 m 16 mia 28 mia 16
mea 48 mia 4 m 440 mia 4O mia 20
m 276 Post-3 mia 36 m 80 mia 32 m 48
mia 56 m 4 m 64 m 488 mia 28 m 4
mia 16 mia 16 mia 4 mia 108 m 4
mia 32 mia 16 mea 36 Post-1 mia 48 mia 8
mia 24 mia 20 m 40 mia 8 m 28
m 36 mia 20 Pre-2 m 4 mea 4 mia 4

92

Table 5 (cont'd.)

 

 

S and S and S and S and S and S and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 36 m 368 m 20 mia 16 Post-1 Post-3
m 184 m 4 mia 12 mia 24 mia 8 m 4
mia 4 mia 12 m 28 mia 28 mia 132 m 4
m 64 mia 8 mia 28 mia 32 m 24 m 4
mia 4 mia 20 m 24 m 40 m 40 m 72
mia 32 mia 20 m 16 mia 20 mia 4 mia 28
m 100 mia 20 m 8 mia 72 m 64 m 48
m 4 mia 20 m 12 mia 4 mia 72
mia 8 mia 24 mia 36 mia 112 mia 16
16 mea 16 neo 36 mia 16 m 68 mia 40
356 m 400 Rep. 2 m 16 mia 56 m 72
84 m 32 Pre-1 mia 28 mia 64 mia 36
24 m 164 m 16 mia 112 mia 48
72 Post-3 mia 60 mia 4O mia 56 mia 28
mia 4 mia 4 mia 76 mia 20 mia 88 m 40
mia 32 mia 12 mia 100 m 20 mia 52 m 36
m 20 mia 24 mia 6O mia 36 mia 64 m 24
m 20 mia 20 mia 24 mea 20 m 72 mia 4
mia 16 m 92 mia 92 mia 36
Post-1 m 28 mia 24 Pre-3 mia 52
m 136 mia 20 mia 52 m 8 Post-2 mia 32
m 4 mia 4O mia 20 m 8 m 12 mia 32
m 40 mia 16 mia 76 mia 20 mia 4 m 28
m 120 m 20 m 16 mia 28 m 8 m 44
m 656 mia 24 m 16 mia 40 mia 4 mia 24
m 112 mia 20 mia 4 mia 32 mia 36 mia 44
mia 24 m 56 mia 32 m 48 m 56
Post-2 mia 20 m 4 mia 28 m 24 mia 12
m 12 m 36 mea 4 mia 56 mia 4 mia 68
m 4 mea 4 mia 20 mia 88 mia 36
m 4 m 324 Pre-Z mia 48 mia 52 mia 52
mia 4 m 16 mia 8 m 28 mia 36 mia 32
m 8 m 116 m 16 mia 44 mia 52 mia 32
mia 8 mia 4 mia 20 mea 20 m 48 m 16
mia 8 mia 20 mia 32 m 372 mia 48 mea 8
m 32 mia 36 mia 36 m 4 mia 56
mia 4 mia 28 mia 16 mia 72 mia 44
mia 12 mia 20 mia 76 m 8 mia 6O neo 4O
mia 16 mia 24 mia 20 m 28 m 28 Rep. 2
m 20 mia 20 mia 36 mia 36 m 48 Pre-l
mia 4 mia 20 m 32 mia 28 m 40 mia 168
mia 16 mia 16 mia 8 mia 28 mia 28 m 28
m '32 mia 20 mia 40 m 28 m 56 mia 20
m 16 m 36 mia 24 mia 44 mea 28 mia 16
m 12 mia 4 mia 24 mia 56 m 240 mia 20
mea 8 mia 24 m 32 m 32 m 20

93

Table 5 (cont'd.)

 

 

g and g and g and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 40 m 56 mia 20 m 36 mia 40 mia 32
m 16 mia 4 mea 20 m 40 mia 8 mia 24
mia 8 mia 28 m 252 m 24 m 36 mia 28
m 36 mia 60 m 24 m 16 mia 8 m 40
m 16 mia 20 m 64 m 4 mia 20 m 16
mia 20 mia 28 m 24 m 80 mia 72 mia 28
mea 24 mia 24 mia 20 m 52 mia 20
m 316 mia 24 mi 12 Post-3 mea 24 m 24
mia 36 mia 24 m 4 m 4 mia 440 mia 16
m 28 mia 44 mia 16 mia 4 mia 36 mia 84
mia 28 m 48 mia 16 mia 12 m 84 mia 20
mia 56 m 20 mia 20 m 16 mia 4 m 24
m 24 mia 16 mia 16 mia 4 m 68 m 20
mea 8 mea 36 mea 16 m 20 m 24 mia 4
m 420 m 12 mia 16 mia 20
Pre-2 mia 24 Post-2 mia 20 mia 28 m 24
m 8 mia 24 m 12 mia 16 mia 4
mia 4 mia 36 m 32 m 20 Pre-2 mia 24
mia 120 mia 28 m 16 mia 4 m 12 mea 20
mia 132 mia 36 m 8 m 20 mia 12
mia 212 mea 28 m 32 mia 20 m 108 Post-1
mia 32 m 64 mea 12 mia 104 mia 12
mia 12 Post-1 m 12 mia 360 mea 60 m 32
mia 20 m 4 m 4 mia 24 mia 400 mia 24
mia 24 mia 4 m 4 mia 16 mia 144 mia 20
mea 28 mia 12 m 4 m 16 mia 112 mia 24
m 416 mia 20 m 24 mia 12 m 164 mia 32
mia 4 m 16 m 28 m I 24 mea 4 m 36
mia 12 mia 16 m 12 mia 4 mia 28
mia 24 mia 24 m 36 m 16 Pre-3 mia 24
mia 20 m 24 m 24 mia 24 m 12 mia 32
mia 24 mia 24 m 24 mia 16 mia 4 mia 56
mia 52 mia 24 m 76 mea 16 mia 32 mia 20
mea 24 mia 20 m 4 m 412 mia 52 m 20
mia 24 m 24 m 4 mia 24 m 48
Pre-3 mia 24 m 20 mia 48 mia 32
m 8 m 16 m 24 mia 24 mia 20
m 4 mea 12 m 52 neo 41 mia 20 mia 32
m 12 m 184 m 88 Rep. 2 mia 76 m 12
m 36 m 100 m 4 Pre-l mia 20 mia 20
m 4 mia 16 m 64 m 24 mia 32 mia 32
m 32 mia 28 m 24 m 8 mia 24 m 40
mia 24 mia 24 m 4 mia 28 mia 28 m 20
m 4 mia 20 m 80 mia 52 mia 24 mea 4
m 44 mia 16 m 8 mia 24 mia 44 m 348
m 4 mia 20 m 128 m 72 mia 16 mia 12

94

Table 5 (cont'd.)

 

§ and g and § and § and § and § and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 64 mia 28 mia 36 mia 40 Post-3 mia 8
mia 12 m 32 mea 40 mia 20 m mia 20
mia 28 mia 16 m 220 mia 40 m 12 mia 32
m 28 m 72 m 16 m 12 mia 20
mia 52 mia 32 mia 24 m 16 mia 20
mea 36 sham 23 mia 24 m 44 m 8 mia 24
Rep. 2 mia 56 m 24 m 8 mia 24
Post-2 Pre-1 mia 32 mia 24 m 4 mia 16
m 16 m 20 m 28 m 16 m 32 mia 52
m 8 mia 28 m 24 mia 4 m 60 mia 16
m 24 m 36 mia 12 m 36 m 48 mia 20
m 12 mia 12 mia 20 mia 16 m 16 mea 20
m 24 mia 44 m 48 mia 48 m 48
m 4 mia 32 m 4 mia 64 m 4 Pre-2
m 24 mia 40 m 8 m 32 m 144 mia 12
m 80 mia 32 m 4 m 12 m 72 m 12
m 44 mia 40 mia 8 m 28 m 4 mia 8
m 108 mia 36 m 16 mea 4 m 12 mia 20
m 108 mia 36 mea 16 m 200 mia 12
m 64 mia 32 Post-1 m 32 mia 20
m 56 mea 48 Pre-3 m 32 m 64 m 20
m 500 m 292 m 4 m 4 m 28 mia 20
m 24 m 32 mia 20 m 20 m 52 mia 36
m 4 mia 28 m 12 m 4 m 112 mia 24
mia 52 m 4 m 40 m 36 m 20
Post-3 mia 52 m 16 m 44 mia 20
mia 4 mia 60 m 8 m 312 mia 24
mia 12 m 40 m 8 m 16 sham 25 mia 20
m 12 mea 28 m 8 m 552 Rep. 2 mia 16
mia 16 m 8 m 4 Pre-l mia 32
mia 20 Pre-2 m 12 m 4 m 96 m 28
mia 20 m 4 mia 4 m 144 mia 24 mia 20
m 24 mia 8 m 12 mia 28 mia 28
mia 4 mia 12 mia 20 Post-2 mia 4 mia 20
mia 16 mia 16 mia 24 m 8 mia 32 mia 32
mia 24 m 20 mia 24 m 16 mia 16 mia 12
mia 28 mia 16 m 20 m 4 mia 24 mia 24
m 20 m 44 mia 24 m 4 mia 24 m 48
mea 44 mia 20 m 8 m 32 m 24 mia 12
m 392 mia 40 m 4 m 44 mia 24 mia 20
mia 4 mia 44 m 16 m 32 mia 24 mea 12
mia 12 mia 28 mia 20 m 228 mia 20 m 320
mia 24 mia 44 m 24 m 20 mea 28 m 28
mia 36 mia 16 m 28 m 424 m 244 m 4
mia 24 mia 52 mia 8 m 4 m 132 m 56
m 24 mia 36 mia 32 m 228 m 4 mia 4

95

Table 5 (cont'd.)

 

 

g and g and § and S and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 12 m 16 m 16 m 4 m 12 m 12
m 44 mia 12 m 4 mia 4 mia 16 m 4
mia 16 m 20 mia 24 m 12 mia 24
mia 20 m 4 Post-2 mia 28 mia 20 m 32
mia 20 mia 20 m 8 mia 20 mia 16 mia 24
m 28 mia 20 m 4 m 28 mia 20
mea 4 m 12 m 4 mia 8 mia 28 Pre-3
mia 24 m 4 mia 20 mia 20 m 20
Pre-3 mia 20 m 16 mea 24 m 16 m 16
m 12 m 24 m 8 m 304 m 24 m 4
m 20 mia 4 m 4 m 4 m 8
m 4 mia 16 m 12 m 20 m 28
m 32 mia 36 m 56 sham 32 mia 24 m 8
m 16 mia 16 m 72 Rep. 2 mia 24 m 12
m 12 mia 16 m 40 Pre-l m 16 m 68
m 20 mia 20 m 52 m 60 m 24 mia 8
m 72 m 16 m 76 mia 12 mia 24 mia 24
m 16 m 4 m 236 mia 56 mia 44 m 16
m 16 mia 4 m 120 mia 20 m 16 m 12
m 56 mia 24 m 440 mia 36 mia 4O mia 16
m 16 mia 32 m 12 mia 64 m 20 mia 20
m 16 m 12 m 16 m 20 mia 16
m 28 mia 28 Post-3 m 36 mia 28 mia 20
m 16 m 8 m 4 m 32 mia 20 m 20
m 36 m 12 m 4 mia 24 m 24 m 24
m 32 mia 4 mia 4 mia 48 m 20 m 4
m 16 m 24 mia 32 m 56 mia 12 m 40
m 112 mea 4 mia 20 mia 4 mia 28 m 40
m 76 m 212 m 8 mia 28 m 32 mia 36
m 8 m 264 mia 12 m 80 m 4 m 24
m 16 m 4 m 20 m 24 m 24 m 4
m 4 m 4 mia 4 m 28 m 24 mia 4
m 32 m 24 m 16 mia 32 mia 20 mia 28
m 28 m 4 m 20 mia 56 m 28 m 48
m 36 m 4 mia 4 mia 6O mia 8 mia 4
m 4 m 4 m 12 m 20 m 16 mia 24
m 4 mia 36 m 8 m 60 m 16 m 48
m 192 mia 8 mia 20 m 52 m 28 mia 24
m 24 mia 32 m 16 m 36 m 16 mia 52
m 76 m 76 mia 12 m 56 m 16 mea 16
m 52 m 4 mia 16 m 4 mia 20 m 224
m 8 mia 16 mea 4 m 12 m 92
Post-1 m 8 mia 16 m 8 m 60
m 4 m 4 mia 16 Pre-2 m 20 mia 8
mia 4 m 4 mea 12 mia 24 m 40 mia 4O
mia 12 m 8 m 420 mia 12 m 44 mia 12

96

Table 5 (cont'd.)

 

S and g and g and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 24 m 24 m 20 m 36 mia 20 mea 20
m 20 mea 4 mia 20 m 24 m 184
Post-1 m 24 m 176 mia 28 m 24 m 92
m 8 m 48 m 28 mia 48 mia 8 m 4
m 4 mia 4 m 64 m 48 m 16 m 4
mia 4 m 20 m 64 mea 4 m 20 mia 96
m 16 mia 32 m 60 m 160 mia 8 mia 20
m 8 m 24 m 64 m 4 mea 16 mia 20
mia 4 mea 4 m 4 m 4 mia 24
m 12 mea 8 m 60 Pre-2 mia 28
m 16 Post-2 m 40 m 32 m 116 mia 20
mia 16 m 16 m 52 m 16 m 56 mia 4O
mia 8 m 4 m 20 m 60 m 52 m 16
mia 8 m 4 m 4 m 4 m 44 mia 16
m 20 m 4 m 24 m 4 m 4 mia 16
m 12 m 8 m 20 m 24 m 4 mia 20
mia 4 mia 4 mia 4 mia 4 m 28 mia 20
mia 20 mia 20 m 28 m 48 m 148 mia 20
m 20 m 4 m 20 m 4 m 32
m 4 m 4 Post-3 mia 16 m 108 mia 20
m 32 m 8 mia 4 m 20 m 4 mia 32
mia 28 mia 20 m 4 m 32 m 12 m 24
m 20 m 28 m 8 m 36 m 4
m 28 m 4 m 8 m 176 mia 16
mia 4 m 12 mia 4 sham 42 m 8 mia 32
m 16 m 40 m 20 Rep. 2 m 196 m 20
m 24 mia 4 mia 16 Pre-l m 32 mia 4
m 24 m 32 mia 28 m 24 m 20 m 32
m 28 m 36 mia 16 mia 24 m 76 m 4
m 40 mia 4 mia 28 mia 20 mia 4 m 4
mia 40 m 32 m 24 mia 24 mia 4 mea 4
m 16 mia 12 m 20 mia 4O mia 20
m 32 mia 16 mia 28 mia 32 m 40 Post-1
mia 16 m 48 m 12 m 24 mia 4
m 20 m 4 m 24 mia 16 Pre-3 mia 12
mia 28 m 8 m 8 m 60 m 4 m 28
m 20 m 24 m 24 m 4 mia 4 mia 12
m 24 m 16 mia 16 mia 16 m 16 mia 24
m 8 m 4 m 24 mea 28 mia 20 mia 36
m 16 m 12 mia 24 m 400 mia 24 mia 16
mia 48 m 24 m 20 m 64 m 28 mia 24
m 16 m 12 mia 16 mia 16 mia 24 mia 8
m 40 m- 4 mia 24 mia 20 mia 16 m 16
m 20 mia 4 mia 24 m 28 mia 28 mia 24
m 16 m 24 mia 48 m 20 m 32 mia 16
m 48 m 4 mia 16 mia 4 mia 16 m 24

97

Table 5 (cont'd.)

 

 

g and S and g and § and g and § and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 16 m 80 m 128 mia 28 m 16 mia 20
m 16 m 4 m 4 m 16 mia 16 mia 24
mia 20 m 4 mia 8 mia 20 m 20 mia 24
mia 20 m 60 mia 32 mia 88 mia 4 m 48
m 24 m 28 mia 24 mia 16 mia 56 mia 12
m 12 m 4 mia 16 mia 20 mia 12 mia 24
m 16 m 44 m 24 m 72 mia 12 mia 32
mia 12 m 236 mia 24 m 8 mia 12 mia 16
m 16 m 4 mia 28 mia 4 mia 24 mia 36
m 20 m 16 mia 32 mea 24 m 20 m 24
mia 4 m 148 mia 20 m 300 mia 4 mia 32
m 28 m 44 mia 52 m 52 mia 24 m 36
mia 24 m 52 m 20 m 12 m 20
mia 12 m 32 mia 32 Pre-2 mia 24 mia 80
m 20 m 8 mia 16 m 20 m 16 mia 12
mia 20 m 4 m 4 m 36 m 52 m 32
m 20 m 172 m 4 m 12 m 8 m 16
m 16 m 4 m 12 m 4 mia 24
mea 8 Post-3 m 4 m 36 m 52 m 28
m 396 m 4 mea 12 m 8 m 4 mia 4O
mia 8 m 4 mia 32 m 12 mia 52
mia 12 mia 4 mia 68 mia 4 mia 16
m 20 m 12 sham 43 m 16 mia 28 m 12
mia 16 m 4 Rep. 2 mia 8 mia 32 mia 48
m 20 m 4 Pre-l m 8 m 24 mia 16
mia 8 m 12 mia 20 mia 4 mia 28 m 28
mia 16 mia 4 m 12 m 24 mia 12 mia 24
mia 16 mia 24 m 16 mia 20 m 16 mia 36
mia 28 mia 20 m 4 mia 20 mia 24 m 32
m 12 m 20 mia 24 mia 16 m 12 m 20
m 8 mia 20 m 16 mia 48 mea 4 m 36
m 8 mia 20 m 16 mea 32 mia 452 mia 4
m 4 m 16 mia 20 m 224 m 16 m 8
m 16 mia 24 m 40 m 68 m 28 m 4
mia 4 mia 24 m 24 m 96 mia 4 m 4
m 28 mia 52 m 76 mia 16 m 4
Post-2 mia 20 mia 48 m 4 m 20 m 4
m 4 mia 24 mia 16 mia 8 m 20 m 32
mia 4 m 24 m 16 mia 28 mia 4 m 4
m 12 mea 24 mia 32 mia 44 mia 16 m 4
mia 68 m 168 mia 32 mia 60 m 4
m 36 m 44 m 12 mea 4 Post-1 m 28
m 4 m 4 m 4 m 4 mea 44
mia 4 m 4 mia 20 Pre-3 m 4
m 20 m 52 mia 20 mia 8 mia 12 Post-2
m 12 m 76 m 44 mia 28 m 16 m 12

98

Table 5 (cont'd.)

 

g and § and g and g and S and § and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
m 20 mia 8 mia 52 m 280 m 4 mia 20
m 40 mia 16 mea 24 m 96 m 48 m 24
m 4 mia 36 mia 284 m 60 mia 12
m 4 m 16 mia 32 Post-1 m 8 m 20
m 12 m 24 m 16 m 4 m 8 mia 12
m 28 mia 44 mia 36 m 4 m 112 m 24
m 44 mia 28 mia 40 m 4 m 364 mia 8
m 80 m 20 mea 24 m 16 m 64 mia 24
m 48 mia 40 m 32 m 312 m 24
mia 4 mia 24 Pre-2 mia 36 mia 4 mia 4
m 12 mia 16 mia 4 m 48 mia 28 mia 16
mia 4 m 24 mia 24 mia 4 mia 24 m 20
m 56 mia 32 mia 24 m 28 mia 20 m 8
m 52 m 20 mia 16 m 12 mia 24 mia 8
m 4 mia 12 mia 28 mia 12 mia 36 mia 16
mia 4 m 36 mia 28 m 16 mia 24 m 12
mia 24 mia 12 m 44 m 20 m 32 mia 4
mia 16 m 24 mia 4 mia 4 m 4 m 220
mia 40 m 12 mia 36 mia 20 mia 4 m 60
m 12 mea 4 mia 24 mia 32
m 28 m 524 m 16 mia 28 Post-3
m 16 mia 20 mia 4 m 24 m 8 nor 30
m 36 mia 24 mia 48 m 12 mia 8 Rep. 2
m 16 mia 24 mia 24 m 4 mia 8 Pre-l
mia 4 mia 36 mia 20 m 8 m 16 m 16
mia 36 m 32 mea 36 mea 4 m 20 mia 8
mia 16 mia 32 m 320 mia 360 mia 24 m 24
mia 48 m 8 mia 28 mia 24 mia 8
m 12 m 28 m 52 mia 24 mia 12
m 20 nor 26 m 4 mia 8 mia 20 mia 4O
mia 12 Rep. 2 m 40 mia 20 mia 12 mia 28
m 20 Pre-l m 64 mia 40 m 16 mia 36
m 24 m 4 mia 52 m 32 mia 4 mia 20
m 4 mia 52 m 36 m 12 mia 12 m 28
m 20 m 20 mia 152 m 12 mia 20 mia 16
mia 20 mia 8 mia 8 m 8 m 16 mia 24
m 24 mia 24 mia 32 m 24 mia 4 mia 28
m 4 mia 4O mia 60 m 16 m 16 m 20
mea 4 mia 28 mia 16 mea 4 mia 4
m 268 mia 32 Pre-3 m 20 m 144 mia 36
m 4 m 32 m 8 mia 28 m 92 mia 48
mia 24 m 20 m 40 m 12 mia 24
Post-3 mia 36 m 24 m 4 m 4 mea 32
m 4 m 28 m 24 mea 4 m 36 m 388
m 4 mia 12 m 148 mia 64 mia 40
m 4 mia 28 m 424 Post-2 m 8 mia 32

99

Table 5 (cont'd.)

 

 

g and g and S and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 32 mia 48 mea 4 m 88 mia 24 m 16
mia 36 mia 20 m 128 mia 16 m 4
mia 24 m 52 Post-2 mia 4 mia 16 mea 4
mea 24 mia 40 m 4 mia 24 mea 28 mia 376
mia 28 m 32 mia 32 m 340 mia 20
Pre-2 mia 24 m 12 mia 16 m 44 m 16
mia 12 m 40 m 8 m 24 mia 24 m 4
mia 12 mia 4 m 8 mia 4 mia 20 mia 40
m 12 mia 32 mia 12 m 16 mia 12 m 16
mia 12 mia 32 m 44 m 8 mia 56 mia 20
mia 28 m 8 mia 4 mia 4 mia 16 m 20
mia 28 mia 4 m 24 m 36 m 20 m 4
mia 20 mia 48 mia 4 mea 8 mia 12 mea 4
mia 12 mia 8 m 20 mea 16
mia 32 m 44 m 4 m 432 Post-1
mia 28 mea 8 m 36 nor 33 mia 4
m 12 mia 4 Rep.2 Pre-3 mia 8
mia 16 Post-1 mia 24 Pre-l m 16 mia 8
mia 12 m 24 mia 20 m 24 mia 8 mia 12
m 36 mia 4 m 24 m 60 mia 16 m 20
mia 16 m 20 mia 4 m 32 mia 8 m 12
mea 20 mia 20 m 32 mia 16 m 8 m 12
m 492 m 44 m 8 mia 36 mia 8 mia 12
mia 4 mia 12 mia 4 mia 36 m 16 mia 20
mia 20 mia 36 m 28 mia 80 mia 28 mia 4O
mia 68 mia 20 mia 24 mia 4O mia 12 mia 16
mia 16 m 32 mia 32 mia 60 mia 16 mia 16
mia 20 mia 20 m 12 mia 68 mia 4O mia 16
mia 28 mia 36 mia 4 mia 52 m 16 mia 20
mia 48 mia 16 mea 24 mia 8 mia 12 mia 16
mia 20 mia 36 m 644 m 76 m 32 m 16
mia 20 m 32 m 8 mia 48 mia 4 mia 16
mia 28 mea 4 m 36 mia 8 mia 32 m 20
mea 32 m 380 mia 4 mea 52 m 36 mia 44
mia 4 mia 20 m 400 mia 20 m 40
Pre-3 mia 56 m 20 mia 56 m 16 m 16
m 36 m 12 mia 8 m 8 mia 12
mia 8 m 16 Post-3 mia 20 mia 12
m 32 mia 32 mia 4 Pre-2 mia 24 mia 36
mia 24 m 40 m 52 m 4 m 16 m 8
m 32 mia 20 mia 24 mia 4 mia 12 mea 20
m 12 m 16 m 24 m 12 m 16 m 264
mia 28 m 36 mia 28 mia 12 m 4 m 8
m 8 mia 4 mia 32 mia 12 m 24 mia 20
m 72 mia 4 mea 20 mia 12 mia 4 mia 12
mia 8 m 32 m 324 m 12 mia 56 m 12

100

Table 5 (cont'd.)

 

g and g and § and g and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 8 mia 12 mia 24 m 32 Post-1 m 92
m 12 mia 28 mia 72 m 176 m 4 m 68
mia 24 mea 20 m 20 mia 32 m 4 m 140
mia 16 m 152 m 88 m 8 m 108
mia 16 mia 196 m 40 Pre-3 m 4 m 196
mia 16 mia 12 mia 4 m 4 m 8 m 264
m 16 mia 20 m 44 mia 8 m 12
m 32 m 12 m 4 mia 20 mia 4 Post-3
m 8 mia 20 m 12 m 36 m
m 16 mea 20 Pre-2 mia 20 m 24 mia 4
mia 20 m 436 m 4 mia 20 m 44 m 16
mia 8 mia 4 m 4 mia 20 m 8 m 16
m 20 mia 20 m 4 m 24 m 16 mia 20
m 24 mia 16 m 20 mia 32 mia 8 mia 32
mia 4 mia 16 mia 16 mia 24 mia 24 m 24
m 24 mia 16 mia 16 mia 52 mia 24 mia 4
mia 8 mia 16 mia 20 m 16 mia 56 m 24
mia 28 m 20 m 56 mia 28 m 48 m 28
mea 16 mea 4 mia 36 m 36 mia 12 mia 4
m 16 mia 4 mia 32 m 28
Post-2 mia 20 m 36 m 28 m 20
m 12 nor 35 mia 16 m 44 mea 20 mia 16
m 4 Rep. 2 m 40 m 4 m 152 m 36
m 4 Pre-1 mia 20 m 48 m 4 mea 4
m 4 m 8 m 36 mia 104 m 16 m 172
m 40 mia 4 m 16 m 20 m 4 m 24
m 64 m 24 m 8 m 24 m 52 m 92
m 32 mia 4 mia 24 mia 20 mia 4 m 152
m 4 m 32 m 4 m 44 mia 28 m 32
m 116 mia 28 m 4 mia 28 m 52 mia 80
m 4 mia 28 m 4 mia 20 mia 28 mia 20
m 44 mia 32 m 4 mia 28 m 40 mia 24
m 72 m 40 m 8 mia 32 m 12 mia 24
m 4 mia 12 m 40 m 16 m 4 mia 20
m 168 m 32 m 32 m 24 m 28 m 24
m 4 mia 44 mia 4 mia 72 m 16 mia 44
m 216 mea 20 m 12 m 16 mia 4 m 56
m 28 m 88 m 16 m 12 m 32 mia 12
mia 32 mia 4 m 28 m 8 m 40
Post-3 m 20 m 36 mia 40 m 8 mea 4
mia 4 m 32 mia 4 mia 28
mia 12 m 4 mia 60 m 24 Post-2
mia 20 m 40 m 28 m 104 m 4 nor 37
mia 16 m 20 mea 4 mia 4 m 68 Rep. 2
mia 24 m 32 m 284 m 24 m 24 Pre-l
mia 20 m 36 m 32 m 80 m 32

1

01

Table 5 (cont'd.)

 

§ and g and g and § and g and g and
beh. irt beh. irt beh. irt beh. irt beh. irt beh. irt
mia 20 mia 28 m 4 m 4 mia 24 mia 32
m 52 mia 28 mia 56 m 4 m 24 m 20
mia 4 m 36 m 16 m 4 mia 20 m 20
mia 52 m 4 mia 4 mia 8 mia 20 m 4
mia 36 mea 4 mia 64 mia 16 mia 20 m 12
m 64 mia 40 m 32 mia 16 m 4
m 4 Pre-3 mia 28 mia 8 m 24 m 8
mia 4 m 4 mia 4O mia 32 mia 4 m 4
m 36 mia 12 mea 28 mia 24 mia 52 mia 20
mea 28 mia 28 mia 20 m 16 m 4
mia 316 mia 28 Post-2 m 24 mia 12 m 8
mia 12 mia 28 m 4 mia 24 mia 8 m 28
m 60 m 36 m 4 mia 12 m 4 mia 8
mia 32 mia 36 m 4 mia 16 m 4 m 32
mea 6O mia 24 m 8 m 24 m 4 mia 168
mia 28 m 20 m 24 m 8 m 16
Pre-2 mia 36 m 24 m 16 m 20 m 16
m 28 mia 32 m 44 mia 16 mia 4 mia 20
m 20 mia 52 m 16 mia 24 mea 12 m 48
mia 20 m 36 m 4 mia 32 m 344 m 28
mia 16 mea 20 m 4 mia 24 mia 24 mia 4
mia 36 mia 404 m 28 m 28 m 20 m 40
mia 20 mia 52 m 104 m 20 m 8 m 8
m 24 mia 32 m 4 mia 48 m 12 m 4
mia 24 mia 28 m 88 m 12 m 4 m 4
mia 36 mia 44 mia 28 m 16 mia 28 m 4
mia 36 mea 32 m 64 mea 20 m 20 m 4
mia 32 mia 8 m 424 m 16 mia 8
mia 16 Post-1 m 28 m 36 m 4
m 20 m 4 mia 4 m 20 Pre-2 mia 12
mia 16 m 4 m 28 mia 4 m 8 m 8
mia 40 m 4 mia 12 mia 16 m 4 m 20
mia 4 m 4 mia 24 mia 36 mia 4 m 12
mia 20 m 32 m 48 mia 28 m 16 m 16
mia 24 m 20 mia 12 mia 24 mia 16 mia 24
mia 20 m 24 mia 36 mia 28 m 24 m 36
mia 24 m 4 mia 28 mia 16 mia 32 m 4
m 24 m 44 m 32 mia 16 mia 24
m 32 m 4 mia 52 mia 28 m 40
mea 4 mia 4 mia 24 nor 38 mia 12 m 4
m 312 mia 72 mia 24 Rep. 2 m 24 m 4
mia 28 mia 76 mia 28 Pre-l mia 8 m 20
mia 8 mia 76 m 40 m 56 m 20 m 4
mia 24 mia 32 mea 8 m 300 m 12 m 4
mia 28 mea 56 mia 4 m 24 m 4
mia 28 m 352 Post-3 mia 28 mia 16 m 4

102

Table 5 (cont'd.)

 

 

g and g and g and g and § and
beh. irt beh. irt beh. irt beh. irt beh. irt
m 4 m 4 m 24 m 16 mia 8
m 36 mia 4 m 4 m 28 mia 32
m 4 m 12 m 28 m 4 m 32
m 4 m 8 m 12 m 8 m 16
mia 8 mia 12 m 20 m 28 m 16
m 8 m 16 m 52 m 4 mia 16
m 12 mia 12 m 4 m 124 m 48
m 4 m 12 m 8 mia 16
m 20 m 12 m 20 Post-3 m 40
m 4 mia 8 m 12 m 4 mia 4
m 4 mia 36 mea 4 m 4 m 64
m 20 m 188 mia 4 m 8
Pre-3 mia 16 m 48 mia 16 m 20
m 44 m 20 m 12 m 24 m 12
m 16 m 24 m 12 mia 12 mia 12
m 28 m 8 mia 92 m 20 m 44
m 20 m 20 m 4 m 12
m 48 m 4 Post-2 mia 8 m 20
m 56 m 24 m 12 mia 24
m 120 m 8 m 20 m 20
m 24 mia 8 m 4 m 12
m 56 m 28 m 4 mia 16
m 24 m 12 m 28 m 16
m 24 m 8 m 12 m 12
m 16 m 8 m 16 m 8
m 8 m 16 m 8 mia 4
m 4 m 4 m 4 m 24
m 20 m 24 m 28 mia 4
m 140 m 16 m 4 mia 32
m 32 m 16 m 28 m 16
m 4 m 16 m 4 m 12
m 40 m 12 m 4 m 4
m 44 m 4 m 36 m 8
m 4 m 20 m 44 m 8
m 16 m 28 m 4 m 4
m 8 m 24 m 72 m 8
m 8 m 20 m 4 m 4
m 16 m 4 m 28 m 8
m 24 m 4 m 4 mea 12
m 8 m 4 m 16 m 284
m 40 m 8 m 44 m 16
m 24 m 24 m 160 m 44
m 248 m 4 m 84 m 32
m 8 m 12 m 148 mia 4
m 32 m 16 m 20
Post-1 mia 20 m 12 m 20

APPENDIX C

APPENDIX C

Table 6

Raw data: postejaculatory responses not occurring within
20 minute sessions

 

 

Subject Rep. Session ReSponse Interval in seconds
cing 2 1 Pre-2 m 236
Pre-3 m 304
Post-3 m 248
cing 6 1 Pre-2 mia 352
Post-1 m 144
Post-3 m 372
cing 7 l Pre-l mia 484
Pre-3 m 500
Post-1 m 292
Post-3 m 452
cing 12 1 Pre-1 mia 104
Post-2 m 328
cing 18 1 Pre-l mia 232
neo 9 l Post-1 m 404
neo l9 1 Pre-l mia 400
sham 1 l Pre-l mia 516
sham 3 1 Pre-l mia 536
Post-1 m 460
Post-2 m 260
Post-3 m 236
sham 10 l Post-3 m 432
sham 22 l Pre-2 m 352
Post-2 m 400
nor 4 1 Pre-i mia 396
nor 8 l Pre-3 m 324
nor 11 l Pre-3 m 372
Post-2 m 264
Post-3 m 304
nor l3 1 Pre-1 mia 412
Pre-3 m 208
Post-1 m 416
nor 15 l Pre-3 m 300
Post-1 m 352
Post-3 m 392
nor 1? 1 Post-1 m 264

103

104

Table 6 (cont'd.)

Subject Rep. Session Response Interval in seconds
Post-2 m 304
cing 24 2 Pre-2 mia 420
Pre-3 m 356
cing 28 2 Pre-2 m 412
Post-2 m 304
cing 29 2 Pre-3 m 472
Post-3 m 276
cing 34 2 Pre-l m 300
cing 39 2 Pre-2 mia 484
Pre-3 m 460
Post-2 m 184
Post-3 m 392
neo 27 2 Pre-l m 572
Pre-2 m 348
neo 31 2 Pre-l m 304
neo 36 2 Pre-l m 344
Pre-Z m 364
Post-3 m 248
neo 4O 2 Pre-l m 320
Pre-3 m 288
Post-1 m 280
neo 41 2 Pre-2 m 480
Pre-3 m 280
Post-2 m 236
sham 23 2 Pre-l m 260
Pre-2 m 156
Pre-3 mia 388
sham 25 2 Pre-l m 224
Pre-2 m 164
sham 32 2 Pre-l m 442
Post-1 m 228
sham 42 2 Pre-l m 320
Pre-3 m 260
Post-1 m 232
sham 43 2 Pre-2 m 412
Post-1 m 132
nor 26 2 Pre-l m 368
Post-1 m 364
nor 3O 2 Pre-l m 392
Pre-Z m 392
Pre-3 m 560
Post-1 m 404
Post-3 m 472
nor 33 2 Pre-3 m 216
Post-1 m 268
Post-3 m 140
nor 35 2 Post-3 m 268

 

 

Subject

nor 37

Rep.

 

2

105

Table 6 (cont'd.)

Session Response

Pre-l mia
Pre-2 m
Pre-3 mia
Post-1 m
Post-2 m

Interval in seconds

 

428
564
552
344
456

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