SOME ASPECTS OF SEXUAL BEHAVIOR

IN DOMESTIC SHEEP, OVIS ARIES

by

EDWIN M. BANKS1)

(Department of Animal Science & Division of General Studies,

University of Illinois, Champaign, U.S.A.)

(With 7 Figures)
(Rec. 25-VIII-1963)

I. DESCRIPTION OF MOTOR PATTERNS

(A) INTRODUCTION

The first extensive attempts to study the biology of reproduction in sheep

were those of McKENZIE & PHILLIPS (1930) and later of MCKENZIE &
TERRILL (1937), in both of which ovarian activity and sexual receptivity
were analyzed. Many more recent studies have extended our understanding
of the physiological, seasonal and breed characteristics of reproduction in
this species and have been summarized by ROBINSON ( I959). Studies of
sexual behavior in sheep that focus upon such statistical parameters as fre-
quency of mating by rams in a competitive situation, the influence of do-
minance, age, and number of rams in a breeding pen have been reported
(HAFEZ & SCOTT, 1962; HULET et al., 1962a, b & c). However, a close
examination of the motor patterns displayed by both sexes during court-
ship has been lacking. One aim of this paper is to present a description of
of male-female interactions culminating in copulation. In the absence of a
basic catalogue of the behavior repertoire leading to successful mating, ana-
lytical procedures attempting to measure the bio-psychological components

1) Present address: Department of Zoology, University of Toronto, Toronto 5,

(Ontario), Canada.
2) It is a pleasure to acknowledge the helpful advice and criticism of Drs A. V.
NALBANDOV and W. H. JOHNSON during the early stages of this study. The following
aided in the collection of data: J. HENRY,E. WILLIAMS, E. HAMLIN,and W. GIBSON.
Line drawings made from single frames of 16 mm film were done by Mrs K. THOMAS.
The Zusammenfassung was prepared by Dr H. FLÜGEL.This study was supported by
research grant MHO3203 from the N.I.M.H., National Institutes of Health.
250

of this behavior would be premature. Presented in the second section of this

paper are some observations dealing with the ontogeny of male sexual beha-
vior. The final section consists of a study of the effects of castration and
subsequent hormone therapy on the manifestations of courtship in the ram.
Although the last two sections are based upon relatively modest numbers of
animals, the results are presented because of their relation to the descriptions
of mating behavior and because the author is no longer in a position to ex-
pand these studies.

(B). METHODS
The descriptions presented below represent composites based upon obser-
vations made over three consecutive annual breeding seasons. In central
Illinois, the annual breeding season extends from mid-August to February
or early March. The flock was composed primarily of Western cross-bred
white-or-black-faced ewes, and in addition, contained some full-bred Targhee
females. The males were of the Southdown, Hampshire, and Rambouillet
breeds. All rams used in this study were vasectomized and were provided
with either a harness to which a marking crayon was attached, or were
painted daily on the brisket area with an oily pigmented dye. Receptive fe-
males were thereby marked on the rear quarters when mounted by the ram.
This technique is commonly used to establish cyclical episodes of estrus in
this species.
Observations were conducted under the following types of conditions.
Preliminary data were collected with the flock composed of Ioo-I2? ewes
and 4-6 rams freely moving about on pasture. During the second and third
breeding seasons of the study a fenced paddock measuring roughly 20' x 70'
was used. An observation booth was installed at one end of the paddock
primarily to afford cover for the observer during inclement weather; it also
reduced distractions which might have interfered with the normal behavior
of the animals, particularly when the group was being filmed.
The observation paddock was supplied with a changing population of ewes
from the general flock in the following manner. Within a month after the
onset of each breeding season, our records of estrous periods of each ewe
enabled us to select females for the paddock. With few exceptions, the non-
pregnant ewe manifests estrus every i5-r7 days during the breeding season.
Females were placed into the paddock one day prior to their anticipated
estrus and remained there until estrus had terminated; they were then re-
turned to the main flock. In this way, a steady supply of females was avail-
able for making observations of mating behavior. One to five rams, kept in
a small adjoining paddock, were released either singly or in pairs during ob-
 251

servation periods. It was assumed that each ram was highly motivated to
engage in courtship behavior for it was only during observation periods
that these particular males had contact with females.
In addition, a series of observations was conducted on a flock of six
ewes and one ram housed in an indoor pen measuring 14' x 20'. This series
differed also in that the flock was subjected to 24-hour surveillance over
two cycles. The procedure used will be further amplified in a later section.
Recording of the observations was accomplished by a number of means,
e.g. written descriptions, voice descriptions made with a portable tape re-
corder, cinematographic recording and finally, a 20-pen Esterline-Angus
operation event recorder activated by a push-button keyboard was used.

(C). MALE MOTOR PATTERNS

During preliminary phases of courtship, the ram displays a number of

Fig. i a, b, c. Nosing the perineum. (All line drawings made from single frames of
16 mm cinematographic records).
252

Fig. 2 a, b, c, d, e. Sequence illustrates display of nudging. Note partial erection

which sometimes occurs with nudging.

discrete motor acts. Individual differences in the tendency to execute given

components and variations in courtship displays of the same individual
from day to day and from one stimulus ewe to another were encountered.
For this reason, the male program, as described below, should be construed
as a generalized scheme subject to individual idiosyncrasies.
When presented with an array of potential mates, the ram has the task
 253

of selecting those in the estrous condition. All of the components of the

male display program utilized in the selection process may be considered
under the general rubric of appetitive behavior, defined here simply as that
behavior bringing about the selection of a receptive ewe.
The first male display in a sexual interaction, nosing (naso-perineal),
appears to afford olfactory and perhaps gustatory information to the ram.
In this act, the ram pushes its face into the perineum of the ewe and, may
also on occasion nibble the external genitalia (Fig. I a, b & c). Another
suggested result of nosing is the possibility that temperature receptors on
the lips of the male may be sensitive to surface temperature changes occur-
ring on the vulva. That there is in fact a cyclical change in the temperature
of the external genitalia has not yet been demonstrated.
The subsequent maneuver in the courtship sequence, the nudge, is a
complex of several subacts. In its typical manifestation, the male orients
behind the ewe, its body pointed in the same direction as that of the female.
Its shoulder, either right or left, is brought into contact with the right or
left flank of the female. One or the other foreleg is extended and then flexed
in a short choppy kicking motion. Simultaneously the ram lowers and tilts
its head sideways and may utter a series of lowpitched vocalizations. The
latter are invariably accompanied by an extension and retraction of the
tongue. The nudge may be repeated several times or displayed only once.
Throughout its execution, the ram fixates the ewe's head. Physical contact
with the ewe's flank is not essential, for the nudge may be displayed when
the female is a foot or two away from the ram (Fig. 2 a-e). Variants of
the complete action pattern are frequently observed. The ram may display
all subacts except the foreleg extension. An extremely abbreviated version
is sometimes seen in which the male approaches the front of the ewe, tilts
his head and utters the courtship call.
One of the more stereotyped acts of the male is evoked by bladder evacua-
tion of the ewe when courtship commences, or during its course. If properly
oriented, the ram attempts to taste the urine while it is being eliminated
(Fig. 3 a-c). Following micturation by the ewe, the ram may lick and/or
nose the moistened ground. It then assumes the posture and facial expression
termed the "Flehmen" by SCHNEIDER (rg3o). During this display, the ram's
head is raised to about a 30 degree angle, its external nares are drawn back
in a flared position, and the upper lip is curled back to reveal the toothless
portion of the upper jaw (Fig. 3 d, e). The Flelm¡,en posture may be held
only momentarily or it may persist for up to two minutes. The role played
by this act is not at all clear. There is considerable individual variation in
254

I
Fig. 3. Sequence illustrates urination by ewe, a; ram nosing urine on ground, b, c, d;
Flehmen, e.

its expression, some rams display it rarely, whereas others do so consistently.

We have been unable to determine whether this rigorously stereotyped act
possesses any signal value to the female, or to other males. SCHNEIDER
 255

(i93o)observed in a variety of artiodactyls confined in a zoo that this facial

expression could be evoked by presenting the subjects with bread or cotton
soaked in several different volatile substances (ether, valerian). We have
corroborated this finding for domestic sheep, using ether-soaked cotton balls.
The act is thus stimulated by a variety of odorants and although it may
have had its origin as a strictly olfacto-sexual response enhancing the male's
ability to detect an estrous female, it is our impression that it does not now
fulfill this function.
Proper behavioral feedback from the ewe, to be described below, leads
to the further phases of the program, starting with the 1nount ¿Fig. 4)

Fig. 4. Tllustrates phase of mounting and erection which typically occurs during
mounting.

during which the ram straddles the rear quarters of the ewe with his fore-
legs. As it mounts, penile erection occurs and is accompanied by pelvic oscil-
lations. The latter continue after I>itro»iissioJ1 has been accomplished. Eja-
culation may be detected behaviorally by an especially deep thrust and is
always followed by dismounting. In our observations, the male did not in-
variably ejaculate prior to dismounting. In many interactions the ram re-
mounted the same ewe up to five consecutive times, engaging in pelvic oscil-
lations during each episode, before behavioral evidence of ejaculation was
manifested.
There were several commonly observed after-reactions on the part of the
ram. It might pause briefly after ejaculating and dismount, and then resume
courtship with the same ewe by nudging or nosing the perineum. The ram
might leave to court another female, or more simply, it might stand quietly
after the stimulus female had removed itself from the immediate vicinity.
These variations were a function of a combination of circumstances,
especially the previous activity of the ram and the availability of other
partners in close proximity.
256

(D). FEMALE MOTOR PATTERNS

A description of the ewe's estrous behavior requires some mention of the
changes occurring in the underlying hormonal support system. Sheep are
classified reproductively as a seasonally polyestrous species. In the geographic
area where this study was made, the breeding season typically begins in mid-
August and terminates from February to early March. During the course
of the season the non-pregnant female undergoes cyclical ovarian activity
culminating in ovulation at intervals of 15-17 days, on the average. Sexual
receptivity, i.e. estrus, is restricted to a period of time ranging from 15-28
hours during this cycle in the ewes studied here. This range is about average
for the species, but it should be noted that breed differences are reported
in this regard, and that such differences also occur in the total length of the
breeding season (HAFEZ, 1952; ROBINSON, 1959).
The display of an anestrous ewe to the appetitive behavior of a ram is,
in essence, either a passive type of apathy if the ram does not persist in
courting it, or an active avoidance of a vigorously courting male. The latter
circumstance was infrequently encountered in our study, and this may very
well be interpreted as an indication of the acuity possessed by males in their
ability to discriminate between the estrous and the anestrous ewe. The evolu-
tion of the communication system between male and female appears to have
reduced to a minimum dysgenic expenditures of energy by both sexes.
The reaction pattern of the ewe on heat varies with ovarian condition.
During the 15-28 hours of estrus, behavioral manifestations range through
a spectrum beginning with low intensity responses through a phase of high
intensity responses and ending with another low intensity phase grading
into the typical indifference of the anestrous female. The temporal sequence
of these phases obtaining in the population tended to be similar for all ewes,
although some exceptions were encountered. Roughly, the first 3-5 hours
of estrus were marked by the following motor patterns, classed here as
low intensity acts. The ewe stands still when approached by the male, head
position depressed and pinnae of the ears somewhat flattened on the head.
As the ram displays, the ewe may swing her head to the appropriate side
and look at him (Figs 2a-e). She may remain standing while the ram nudges,
or the female may walk of a few steps and stand, again swinging her head
back to look at the ram. If the latter is particularly persistent, the ewe will
accept service.
The first phase of low intensity responsiveness is supplanted by a period
of time during which the ewe is notably more aggressive when approached
by a courting ram. When in this phase of estrus, the ewe may actually
initiate the sexual episode by approaching the male, nuzzling his forequarters.
 257

To this soliciting behavior, the ram's typical response is one of vigorous

nudging. It is interesting to note that in such a situation, the male spends
little time in nosing the perineal region of the ewe. In its most extreme
expression, the ewe pushes her head into the ram's flank, her nose directed
to the scrotal area. This act is accompanied by gentle pushing movements
against the ram. The latter responds by movements which once more orient
him properly to achieve mounting, only to have the ewe wheel around and
again place her head close to the scrotal area. These reciprocally stimulated
actions result in a pirouetting or circling phenomenon, and appear to bring
about an enhancement of the ram's attempts to mount. This high intensity
soliciting was observed between the 5th and r 5th hours following the onset
of estrus.
Low intensity soliciting, as previously described, supervenes until the
end of the estrous period (i5-28 hours), after which time the ewe excites
little interest in the ram.
Ancillary observations revealed that ewes exhibiting low intensity soli-
citing rarely attempted to interfere in the courtship interactions of a ram
and another ewe. Contrariwise, high intensity phase ewes persistently
attempted to interfere when the ram was attending to another ewe. It has
been observed that high intensity ewes can completely disrupt ongoing sexual
interactions. Anestrous ewes were never observed to exhibit this type of
interference. Further observations on interference in sexual activity by the
female are noted below.

(E). OBSERVATIONS ON CLOSED FLOCK

On the basis of past estrous performance, two groups of three ewes
were housed in a small (14' x 20') indoor pen. The anticipated estrous
periods of the two groups were spaced one week apart. A single, sexually
experienced, vasectomized ram completed the pen population. Twelve hours
prior to the expected time of estrus for each group of three ewes, a 24-hour
surveillance of the pen was instituted. Observations were continued until
after all ewes of each group had completed their respective estrous periods,
an interval ranging from r5-28 hours each in these females. In one corner
of the pen, the observers were housed inside a 6' x 6' booth that was
provided with two small observation windows. Light was provided at night
by two 24-watt bulbs fitted in ceiling fixtures.
In addition to data relevant to sexual behavior summarized in the pre-
ceding section, an opportunity was provided by this setting to observe certain
other aspects of social behavior. Although agonistic encounters among rams,
leading to the establishment of dominance are well known, such social or-
258

ganization among ewes has not been reported. Episodes of bunting and sham
fighting occur among ewes in a flock on pasture, but such displays are rare
enough to have apparently discouraged study. In the small indoor pen, con-
ditions were ideal for the determination of dominance relationships, should
such in fact exist. Observations recorded over four sessions of 24
hour surveillance revealed a stable, linear dominance order among the ewes.
Agonistic behavior consisted of head to head bunting, and bunts directed to
other body regions. The submissive component was manifested exclusively
by yielding or avoidance by the subordinate ewe.
It soon became evident that each ewe tended to occupy a particular site
on the pen floor when lying down for rumination, resting, or for sleeping.
That these sites constituted a defended area and, hence, a territory, in the
widest sense, was evidenced by observations such as the following. When
the rumination site of a higher ranking ewe was temporarily vacated, as
would occur if it went to the water tank, a lower ranking female might settle
down in it. Upon returning to its site, the dominant ewe persistently kicked
its flock mate until the latter stood up and moved away. The converse was
never observed, i.e. sites of lower ranking ewes were never usurped by
dominants. Inasmuch as the floor space was quite uniformly covered with
bedding straw, the features used for site selection were not evident to the
observers. It should be noted that a breakdown in this pattern occurred when
the ewes were experiencing estrus. The tendency then was for the females
to settle down close to the ram. The counterpart of this behavior has been
reported frequently in studies made on free-moving flocks in a pasture
situation (INKSTER, 1957). Here, estrous ewes tend to follow the ram.
The sexual behavior observed in the indoor pen was similar in form to
that previously described. It was possible to add to this picture certain aspects
not readily seen in the outdoor paddock or in the pasture observations. The
most significant addition pertains to the role of dominance in the manifesta-
tion of estrus. As noted before, the ewes had been selected so that three
would be in estrus at approximately the same time. With but one ram avail-
able, it became clear that there was a considerable amount of competition
among those females whose estrous periods overlapped. This competition
took the form of physical interference of ongoing courtship between the ram
and one of the ewes. Its most typical manifestation may be described as
follows.
One of the three ewes entered estrus and was courted and mated several
times. Toward the end of its estrous period, a second ewe entered its estrous
period. It was most often observed that the ram would cease courting the
first female and direct his behavior to the second. The ewe first in heat
 259

attempted to insinuate itself between the ram and the female being courted.
This was accomplished by bunting the flank of the other ewe while the ram
courted it. In most instances the ram persisted in his courtship displays
toward the second ewe and eventually achieved a mating. During the four
24-hour surveillance periods, interference of this type resulted in a female
remaining unmated on four occasions. In each of these instances, the inter-
fering ewe held either alpha or beta rank in the linear dominance order
established among the six ewes.
Granted that the living circumstances within the confined space afforded
by the indoor pen could be construed as quite "unnatural", it nonetheless
was established that low-ranking ewes might not be mated, despite their
receptive condition, because of interference provided by estrous females
higher in the dominance order (c f . HULET et al., y62c) . It should be noted
that behavior disruptive of courtship was displayed only by ewes in estrus.
Females not in heat appeared to disregard the courtship activities going on
about them.
Another observation of interest related to the apparent capacity of the
ram to remember the individual with whom it had copulated some hours
previously. For example, after courting and achieving intromission and eja-
culation with a given female several times in succession, the ram might not
have an interaction with this female for a period of up to five hours. In
several such instances, the ram gave every appearance of remembering the
identity of this female. Indication for such memory was best evidenced
when the ewe was lying down, ruminating or resting. The ram was observed
to approach her, to kick her persistently until she stood up, and then to
mount her with no preliminary courtship whatever. Although this sort of
procedure was observed only six times during the four 24-hour surveillance
sessions, the seemingly deliberate selection of a recumbent estrous female in
the manner described was considered ample evidence of memory for indivi-
duals by the ram. During the period intervening between a given sexual
encounter between ram and ewe, the former was, of course, involved in
courtship activity with at least two other females currently in estrus. At no
time was the ram observed to kick a female lying down that was not in an
estrous condition.

(F). TIMING AND SEQUENCE OF ACTION PATTERNS

Observation of courtship patterns among adult, sexually experienced
animals gave the impression of smoothly coordinated sequences of actions
and reactions. However, an examination of filmed records with a time and
motion projector revealed that the several components in a bout of male
260

courtship behavior could, under no circumstance, be described as rigidly

timed acts. There was wide variation in the total time per bout as well as
in the time required for the display of individual acts. Two features only
of the entire male program did provide some indications of time and form
stereotypy, e.g. the Flehmen response and the time required for the execu-
tion of pelvic oscillations. As noted previously, the Flehmen could be held
for as long a period as 2 minutes, but in measurements made of 12 filmed
interactions the average time was of a much shorter duration, 7.6 sec. (31.4
to 2.6 sec.). GEIST (ig63) gave the average of five timed Flehmen respon-
ses in the moose (Alces a. andersoni) as 17 sec. and noted that the duration
was variable in this species. In the 12 filmed interactions, the number of
pelvis oscillations per intromission varied from 3 to 8 and each oscillation
required from 0.5 to o.9 sec. for execution.
One of the interesting problems associated with descriptions of social in-
teractions between two conspecifics is that relating to the phasing of acts
in a bout of species-typical behavior. The ethological literature dealing with
fixed action patterns of social behavior among fishes and birds, in the verte-
brate series, supports, in general, the concept of the importance not only of
the duration of an act in a sequence, but also of interact timing (NELSON,
1962). Accordingly, an attempt was made to record interactions between
males and estrous females in such a way that an evaluation of the import of
time scores and frequencies could be made.
Each act of the male and female repertoire was coded on a 20-key key-
board which activated an operation event recorder. During an interaction,
both frequency and duration of the components of courtship, as described
previously, were recorded from approximately ioo episodes. These data
were then transferred to I.B.M. cards for purposes of tabulation and
analysis.
Because the role played by the male was so very much more active and
evident than that of the female, it seemed reasonable to assume that the
behavior of the male exerted control over the outcome of an interaction.
However, no clear-cut trends were established between the frequency and
duration with which a given act was displayed by the ram and the response
of the ewe except as noted below. For example, whether the ram displayed a
burst of 5 nudges in rapid sequence over 2 sec., or displayed the 5 nudges in
a more deliberate manner over a 25 sec. interval had little or no effect on
the response by the estrous ewe. This held generally for all male precopu-
latory acts. On the other hand, the overwhelming importance of the estrous
condition of the ewe, as reflected in her willingness to stand for mounting
was strikingly illustrated by this analysis. Thus a ewe at the peak of recep-
 261

tivity might accept the ram after only an abbreviated display of courtship,
whereas the same ewe when in the low intensity phase and when approaching
anestrus would tend to be almost completely refractory to the ram. Under
such a circumstance, the ram might achieve copulation only after manifesting
the most strenuous efforts. Hence a relation between high frequency nudging
and acceptance by the ewe would occur. Most rams tended to leave such
refractory ewes and to address their courtship to other prospective mating
partners present in the paddock. The ram was actually much less "in charge"
of the proceedings than one would have assumed on the basis of the more
active role played by the male in sexual encounters. It was only during the
earliest and latest phases of estrus that such trends associating male and
female activity could be detected.
It is, perhaps, not too surprising to find a lack of fixity in the expression
of the action patterns involved in courtship in this species. It may be argued
that the process of artificial selection with strong pressure being exerted
for high reproductive capacities has tended to amputate or blunt the some-
times subtle kinds of signals reported in the more fixed reaction systems
found among many feral species of vertebrates. At the same time, the
description of the display components presented above is representative and
can certainly be used by the casual observer to identify a heterosexual en-
counter among sheep as being sexual in nature.

II. THE ONTOGENESIS OF MALE MOTOR PATTERNS

(A). INTRODUCTION
This phase of the study was designed to obtain insight into the following
questions: I) Is there a chronological sequence of expression of the com-
ponents ? 2) Is the development of male sexual behavior contingent upon
the presence of gonadal hormone? 3) What will be the response of ram
lambs, isolated from heterosexual contact between weaning and well past
the age of physiological sexual maturity when first exposed to a receptive
ewe?

(B). METHODS
This section of the report is based upon a study of eight ram lambs, four
of the Hampshire breed acquired from a commercial breeder and four of
the Rambouillet breed from the University of Illinois flocks. The animals
were housed in the indoor pen described previously, and in addition had
access to an outdoor paddock ioo' x 6o". The lambs ranged in age from 79
262

to 136 days when the flock was formed in March, 1962, all having been
weaned at 60 to 7o days and maintained in unisexual male flocks until used
in the study.
Systematic observations were made between March, ig6i and February,
1962 on the general behavioral interactions of the animals. It will be noted
here that complete assimilation to form a single, cohesive flock required
about iy2 months of living together. Until that time, the four ram lambs of
each breed clustered together. More will be said concerning the types of so-
cial interactions observed at a later time.
One ram of each breed was castrated upon arrival, the Rambouillet (# 13)
at 118 and the Hampshire (# 18) at 139 days of age. One ram of each
breed was deprived of heterosexual experience until after the onset of
physiological sexual maturity, determined by others on the basis of testes
weight and rupture of the preputial membranes to occur on the average at
210 days of age (RowsoN, 1959). At weekly or biweekly intervals all but
the female-deprived rams were tested in the following manner. The entire
flock of eight rams was driven out of the pen into the adjoining paddock.
During the breeding season, a ewe known to be in estrus was introduced
into the ram's pen. Tests were also conducted outside the breeding season,
during the summer of ig6i, in which case injections of progesterone and
estrogen were used to induce the estrous condition in the stimulus animal. In
either event, the stimulus ewe was exposed to an adult, vasectomized teaser
ram prior to testing to assure that it was, in fact, in heat.
After introduction of the stimulus ewe into the ram's pen, each male with
the exception of the two female-deprived rams noted above, was allowed a
lo-minute period with the female, during which time its behavior was
recorded. After having reached 404 and q.I9 days of age, respectively, the
two female-deprived rams were also allowed io-minute exposures with the
stimulus female. Observations were made from the small booth described
previously which now was fitted with a one-way mirror.

(C). RESULTS

(I). General I observations s off behavior r during test t

periods conducted between March 1061 1 and February
IC)62.
During the initial test exposures to the stimulus female, the rams all
behaved in a hyperexcited manner. The two breeds differed strikingly in
their adaptation to the test situation. Rambouillet rams were, by far, more
excitable as reflected in pacing about the pen, frequent vocalization, and
 263

climbing the walls near the doors to the outside paddock. In contrast, the
Hampshire rams rarely vocalized, never climbed the walls and only occasion-
ally paced about the pen. All rams from the onset of testing engaged in
tactile-olfactory examination of the stimulus female. Nuzzling the coat and
nosing the perineal region were actions displayed by all rams, even during
their initial test with the ewe. Such behavior along with naso-nasal contact
could be interpreted as a greeting ceremony, the result of which could be
the establishment of sensory information leading to individual recognition.
One pattern, not encountered when observing adult, experienced animals,
was the nosing of the udder region of the ewe. It almost appeared as though
the rams were attempting to suckle.
The behavior displayed by the stimulus ewes varied somewhat. Prior to
the introduction of the first ram on any testing day, all ewes were allowed a
io-inintite period to investigate the pen. It was not possible to select the ewes
so that all were in exactly the same phase of the estrous period when used
for testing. This also proved to be the case during the nonbreeding season
when estrus was induced in the stimulus ewes by injections of exogenous
hormones, for individual ewes responded differentially to the hormone treat-
ment. As a result of this variation in the estrous condition of the stimulus
ewes, their reactions to the courtship displays of the rams tended to vary
somewhat, but, as stated previously, all were tried with experienced, vasec-
tomized teaser rams and found to be receptive prior to their use in the test
situation. Variability occurred mainly in the degree to which individual ewes
manifested soliciting behavior.

(2). Specific courtship displays s of unaltered rams.

Noted in Table i is the age at which each unaltered ram executed its first
complete copulation. Others have determined that sexual maturation occurs
somewhat earlier in the Hampshire breed than in the Rambouillet. This fact
is borne out by our data. Notice, however, that none of the unaltered rams
executed a complete copulation on the first exposure. In fact, the ram ex-
hibiting the earliest complete response did so on the tenth exposure. The
Rambouillet ram showing the complete response at the oldest age did so on
the 17th exposure at which time he was 483 days old.
With one exception, the unaltered rams exhibited the nudge during tests
prior to that in which they were observed to execute the complete sexual
response. In one case, ram # 17, the first observed nudge and the first
complete copulatory response occurred during the same test, after the ram
had been exposed to a stimulus ewe for 19 trials. In one case, ram # 20,
 264

TABLE 1
Undtered rams

i ) Rambouillet ; 2) Hampshire.

nudging occurred upon the ist exposure to the female in the test situation.
In summary, none of the unaltered, young rams displayed a complete
copulatory response on the first or even the first few exposures to an estrous
ewe. Indeed, all were well past the age of physiological sexual maturity at
the time of their first intromission and ejaculation.

(3). Specific behavior displayed by castrated rams.

Noted in Table 2 are those data obtained from castrated rams. It will
be seen that the Hampshire ram, castrated at i39 days of age, exhibited the
first nudge and first complete copulatory response at 396 days of age, on
the i2th exposure to a stimulus female. The response was not truly com-
plete in that, owing to an immature penile size, intromission was not achie-
ved. However, this ram mounted in proper orientation, exhibited that degree
of erection of which it was capable and manifested pelvic oscillations. Beha-
vioral evidence of ejaculation was absent. The other castrated ram, # 13,
showed no courtship behavior to stimulus females. This ram was subjected
to castration at an age 21 days younger than the Hampshire ram. Whether
this difference accounts for the variance in observed behavior between the
two animals is problematical.

TABLE 2
Castrate rams

1) Rambouillet; 2) Hampshire; 3) incomplete (see text).

 265

(4) Specific behavior r of f female-deprived rams.

Two rams, # 14 and # 23 were deprived of contact with females from
weaning age (about 7o days) until they were 404 and 419 days of age, res-
pectively. Noted in Table 3 are the pertinent data. The Rambouillet # 14
evidenced no copulation at the time of his 12th exposure, being then 676
days of age. On the other hand, the Hampshire ram # 23 displayed a

TABLE 3
Female-deprived rams

complete sexual response upon its first exposure to a stimulus female. At

this time, it was q.r9 days old. Of the unaltered rams two exhibited a com-
plete reaction to the stimulus ewe at younger ages than did ram # 23. In
one case, # 20 succeeded at 358 days of age, but not until the tenth exposure,
whereas # 2 displayed the complete response at 406 days, on the 15th
exposure to the stimulus female.

(5). General I observations s of f ram flock.

Before attempting an interpretation of the observations presented above
it would be well to discuss the nature of the observed inter-ram social be-
havior. Observations were made at regular intervals of the ram flock when
it was inside the pen and when the animals were moving about the outdoor
paddock. The purpose of these observations was to determine the types of
social behaviors displayed and to see whether any of the components or acts
of adult courtship were displayed in this all-male group. At times, observa-
tions were conducted inside the pen just after a standard ration of mixed
grains had been placed in the feeder. It was hoped that competition for the
feed would enable us to assess the relative level of aggressiveness of each
ram.
It was pointed out previously that for the first iy2 months after the flock
was assembled, the four rams of each breed tended to stay close together,
forming, in effect, two distinct groups. Whatever the barriers were to the
formation of a single flock, these gradually disappeared and by two months
after assemblage, a unitary, cohesive flock had formed.
No effort was made to observe systematically such behaviors as feeding,
 266

drinking, rumination, and elimination. Rather, we were most interested in

noting the kinds of social interactions which characterize males at this age
and in this peculiar environmental circumstance.
During the course of normal movements about the paddock, flock mem-
bers remained close to one another. The most prominent type of inter-indi-
vidual behavior could be operationally classified as investigatory in nature.
Such acts as head-rubbing, nuzzling the coat of a flock mate, nosing the
body and especially the perineal region of a flock mate. A curious activity
which occurred frequently, was that of nosing the region between the hind
legs of a conspecific, oriented as a lamb orients to suckle the ewe. As noted
previously, this area attracted investigatory attention by rams during the
course of their test periods with estrous ewes. There was no way of deter-
mining whether this was simply relict behavior carried over by the ram from
its suckling period, or whether the rams were attracted by the odors emana-
ting from the external genitalia of their flock mates. Frequent bouts of sham
fighting and typical head-to-head bunting were observed.
At rare intervals the flock was observed to engage in gambolling about
the paddock. That such displays were observed very infrequently is pro-
bably a function of the relatively restricted space available, for it is quite
routinely seen among young sheep on pasture.
Of primary interest in the context of the present study, were those dis-
plays consisting of components of adult courtship behavior. Table 4
presents a tabulation of the components of sexual behavior displayed at
least once by each ram during interactions with male flock mates. This
summary includes observations begun when the flock was first assembled
and continued long after all rams had reached physiological maturity.
Note that all unaltered rams displayed the major components from the

TABLE 4

Components af adult courtship displayed by rarns in male unisexual flock

1) Rambouillet ; 2) Hampshire.
 267

onset of our observations. When engaged in these activities, the animals

appeared to be playing. As time passed, the intensity of pelvic oscillations
appeared to increase, but we have no quantitative data to support this im-
pression. It seems to us to be a safe assumption that this play activity had
been engaged in long before we obtained the animals for our study. The
only qualitative difference between this behavior and adult courtship, except
for the absence of the consummatory components, was in the occasional
faulty orientation of the mount, a young ram sometimes mounting a side
or head end of its flock mate.
The two rams deprived of female contact between the time of weaning
and sexual maturity likewise manifested the major acts of adult courtship
in interactions with flock mates.
Of the two wethers, the behavior of the Hampshire # 18, castrated at
139 days was indistinguishable from that of the unaltered rams with the
exception that penile erection was very much less evident. The Rambouil-
let # 13, castrated at i 18 days of age was never observed to mount, achieve
erection, or engage in pelvic oscillations, although it did rarely show nudging.
Nosing of the perineal and belly regions of flock mates was displayed as
frequently by this castrate as by any other member of the group.
Agressive behavior in the form of pushing and shoving at the grain
hopper and frequent short episodes of bunting were observed. It is also
noteworthy that the nudge, a very prominent aspect of adult courtship dis-
play, was also seen when two young rams were engaged in an agonistic
interaction.
Owing perhaps to their faster rate of maturation, all of the Hampshire
rams appeared to dominate the Rambouillets. During the first few months of
study a clearly defined, stable dominance order was not present, there being
many instances of shifting dominance. Competition over the daily ration of
mixed grains was not sufficiently intense to clarify all possible dominance-
subordinance relationships. However, it was clear that the Rambouillet
castrate # 13 held omega status in the flock. Whether its apparently
low level of aggressiveness was in some manner associated with the poor
showing of this ram when tested with estrous ewes is problematical. Of all
the rams, #13 was never observed to mount a flock mate, although he did,
rarely, nudge another ram prior to giving way in an agonistic interaction.

(6). Discussion off ontogeny of f male motor patterns.

It appears that, with one exception, all rams had developed the preliminary
motor patterns used by adults in courtship long before they had reached the
age of physiological sexual maturity. Of the two withers, one was able to
268

execute the pattern of courtship except for true copulation on the 12th test,
whereas the other had not achieved the response.
Of the two rams deprived of heterosexual experience between weaning
age and about ?oo days, one ( # 23) exhibited the complete response on the
first trial (419 days old). The other (# 14), at 676 days of age and on the
12th test had not executed the complete copulatory response.
The response of # 14 to estrous ewes resembled that of # 23 and of the
unaltered ralns in some respects, i.e., with increasing heterosexual experience
this ram manifested enhanced sexual arousal. Ram # 14 differed from all
other unaltered males in that he was never observed to mount and copulate
with a stimulus female, even though all appetitive acts of the male repertoire
were displayed in a normal manner. One possible clue to the inadequacy of
# 14 was his display of an aberrant act termed here "intention-bunting".
In this act, the ram appearcd to be operating under two incompatible drive
systems, namely, agonistic and sexual. The overt symptoms consisted of an
approach to the ewe, followed by a seemingly deliberate "back-up" maneuver.
This was accompanied by a lowering of the head and the observer was led
to expect that a forceful bunt would be delivered to the ewe. In other words,
the male displayed an intention movement to bunt in the ethological sense.
However, the rapid advance to the female ended not with the anticipated
bunt, but rather with an exaggerated nudge just prior to making physical
contact with the ewe. The ewes responded to these displays with some varia-
tion, the major tendency being to orient to the approaching male with head
lowered, as though to receive a bunt in typical fighting fashion. In other
instances, the ram's maneuver simply evoked a rapidly executed avoidance
by the female. In short, this peculiar behavior tended to upset any previously
developing courtship relationship, and would, of itself, have been a sufficient
deterrent to normal sexual interaction that # 14 would not achieve copula-
tion. ,
It remains to add that this ambiguous behavior was not a constant feature
in tests of # 14. In those sessions when intention-bunting was not observed,
# 14 displayed only desultory interest in the stimulus female. It is suggested
that the apparent incapacity of this male was partly the result of the devel-
opment of an aberrant motor pattern. The etiology of this maladaptive be-
havior appears to involve activities concerned with the establishment of
general agonistic responses in the males of this species. This behavior was
observed, rarely, during the course of agonistic encounters among the males
when moving around the outdoor paddock area adjacent to the indoor test
pen. under these conditions, the maneuver had the appearance of a low
intensity agonistic act, displayed by all young rams, except the castrate # 13.
 269

It is by no means clear why only # 14 displayed the intention-bunt in the

test sessions under discussion. Further comment on this display will appear
in a later section of this report when the results of androgen treatment are
presented.
It is most interesting that, with one exception, all animals which did ex-
hibit the complete sequence did so only after from 12 to 20 ten-minute expo-
sures to stimulus females. Estrous ewes did not evoke sexual arousal in
these rams until after they had had considerable experience with them. This
finding could mean that the array of stimuli that evoke the complete response
is, to a substantial degree, learned during a series of initial encounters be-
tween male and female. Another interpretation which focusses attention on
the rearing procedure followed in this study might explain the finding on
the basis of the development of homosexual tendencies (cf. KAGEN &
BEACH, 1953). In its development, the tendency to mount another male, al-
though not reinforced by the achievement of ejaculation may have inter-
fered with the early establishment of the normal heterosexual communica-
tion system that leads to copulation. SCOTT in describing the devel-
opment of social organization in flocks of sheep reared under more normal
conditions than our animals, noted that incipient sexual behavior among ram
lambs was seen as early as day 3 in the form of attempts to mount other
lambs or even adult ewes. This behavior, categorized by SCOTT as play
activity, was quite marked during the fifth and sixth months of life. It
would appear that social interactions of this type occurring early in the life
of the individual and assuming the form of incipient sexual behavior may
be necessary prerequisites to the complete establishment of sexual arousal
and copulation after the male has reached sexual maturity, in this species.
Data from the present experiment offer qualified support for such an argu-
ment. However, one animal, deprived of heterosexual experience until well
past the age of physiological maturity succeeded in displaying the entire ga-
mut of sexuality on the very first exposure to the stimulus female. Another,
of a different breed, but similarly treated, failed though tested until he
was approximately two years old. Within the modest sample of two indivi-
duals, the whole spectrum of possible consequences of the treatment was
expressed. A full-scale examination of this question would be of obvious
value.

III. EFFECTS OF CASTRATION AND HORMONE TREATMENT

(A). INTRODUCTION
Information regarding the persistence of male sexual behavior following
castration is available from numerous studies on rodents, lagomorphs, carni-
270

vores, and primates (YOUNG, 1961). The evidence supports, in general, the
argument that a phylogenetic trend toward increased cortical control of male
mammalian sexual behavior is accompanied by a decreasing dependency upon
hormonal support (BEACH, 1947a, 1947b, 1956). Thus among the lower
mammals, as exemplified by rodents, castration of adult males leads to a
relatively rapid decline in the manifestations of sexual behavior, whereas
similar treatment to primates may result in little or no detectable decrement
in the display of sexual behavior for pcriods of time measured in years. In
a study of this phenomenon in male cats, ROSENBLATT & ARONSON ( i958)
reported data lending support to the importance of experiential factors in
the persistence of post-castrational sexual behavior.

(B). METHODS
In the preceding section, the development of sexual behavior in a small
sample of rams subjected to a variety of treatments was discussed. Obser-
vation of this flock provided data regarding the number and duration of ex-
posures of each ram to estrous ewes and also provided a complete record of
the displays manifested by each ram during test sessions. These data were
recorded during the period of March, 1961 to February, 1962. Three rams
from this group were selected for a study of the influence of androgen
treatment on sexual behavior, viz. # 21, # 18, and # 13.
Ram # 21, (now 478 days old) previously unaltered and displaying con-
sistent adult sexual behavior from the I sth exposure to an estrous ewe was
castrated 26 February, 1962. Rams # 18 and # 13 had been castrated at an
earlier age (Table 2). Whereas # 13 had never displayed sexual behavior
adequate to achieve copulation, the behavior of # 18 was solnewhat puzzling
in that after 11 exposures to an estrous ewe it had accomplished what was
termed incomplete copulation (see pag. 264). As indicated below in more de-
tail, # 18 continued to display this type of behavior during test sessions
until late spring of 1962, at which time all measures of sexual behavior dis-
appeared. Observations recorded from tests of # 21, # iS, and # 13 follow.

(C). RESULTS
(i). Ram # 21.
Attention was focussed upon four major parameters of the male display
pattern, e.g. nosing, nudging, mounting and abortive mounting. Behavioral
evidence of ejaculation was not observed during these tests. An abortive
mount is defined here as a mount terminated prior to intromission as a
result of the activity of the ram, such as an improper orientation. Twenty-
two ten-minute tests were run on all the males between 23 February and
10 August, 1962 at which time testing was discontinued for approximately
 271

two months. The data obtained during these tests from ram # 21 are pre-
sented graphically in Fig. 5. Within a month after castration, the frequency
of mounting by # 2i declined from an average of two per test to zero about
9 weeks after castration. There was a gradual decline in the frequency of
nosing but little emphasis can be placed on this measure because it is the
most generalized type of inter-individual act. In addition to its possible use
by the ram in the detection of an estrous ewe, naso-nasal and naso-perineal
contacts may also provide a means of establishing individual recognition
between two previously unacquainted sheep. No instances of abortive moun-
ting by # 21 were observed during this series of tests.
The testing procedure was reinstated on 3 October, 1962 prior to the
schedule of androgen injections begun on 15 October. Note that the values

Fig. 5. Record of ram # 21, castrated 2/26/62, at 478 days of age.

obtained during the pre-treatment series for frequency of mounting, nudging,
and abortive mounting remained low. Ram # 21 received four weekly in-
jections of 100 mg. testosterone cyclopentylpropionate 1), IM. Four days
after the first injection, # 21 displayed a striking return of sexual behavior.
Maximum activity was evoked by the injected androgen on 30 November,
some twenty days after administration of the fourth and final injection of
the series. There followed an abrupt decline in all measures such that by
8 January, 1963, approximately 2 months after final injection, sexual be-
havior was indistinguishable from the pre-treatment performance.
1) Upjohn Depo-Testosterone.
272

Two facts worth pointing out at this time are ( r ) the frequency of
mounting per io-minute test increased well above pre-treatment measures
(maximum of I5) under androgen support, and (2) abortive mounting, pre-
viously unrecorded for # 21, reached sizeable magnitude.
The inadequacies inherent in using the foregoing method for describing
behavioral data are well documented in this case, for a reading of the protocol
of these observations reveals information that simply cannot be communi-
cated by a presentation of numbers alone. For example, the general aspect of
# 21's deportment, both during tests and outside the test sessions was one
of extreme hyperexcitability. Aggressive behavior in the flock increased
markedly with the androgen support. Most of the abortive mounts were the
result of improper orientation by the ram, including mounting the ewe face-
on, and executing pelvic oscillations while in this position. Such maladroit
activity was never observed in normal males. During test sessions it was
freuqently observed that the ram would back up, approach the ewe with head
lowered, in a manner indistinguishable from that used during bouts of bun-
ting with other males, and then at the last moment execute a nudge instead
of the bunt anticipated by the observer. This action pattern was referred to
previously under the term "intention-bunting". It appeared as though the
threshold for the display of aggression had been so lowered by the injected
hormone, that the stimuli required to evoke bunting operated at a much
lower valence than normal. At the same time, the sexual stimuli emanating
from the estrous ewe were apparently recognized, as indicated in the much
enhanced mounting scores of the treatment period.

(2). Ram # 18.

Comparable data obtained from #18, castrated on 20 March, y6I at the
age of 139 days, are presented in Fig. 6. Intromission disappeared from
the repertoire of this ram early in April, 1962 and remained at zero fre-
quency until tests were terminated in August, rc?62. Nudging continued to
be displayed by # 18 until mid-July, at which time it, too, disappeared.
Nosing, both naso-nasal and naso-perineal continued to be displayed with
varying frequency throughout the tests.
Two tests conducted early in October, after the approximately two-month
hiatus of observation, revealed that the frequency of mounting, nudging, and
abortive mounting continued at zero level. As indicated in Fig. 6, # 18
received the same number and dosage of injections of androgen as had # 21.
Increases in frequency were observed first in nosing, then nudging, and
finally in mounting. The maximum frequency for mounting was recorded
eleven days following the final injection of testosterone. Abortive mounting
 273

Fig. 6. Record of ram #18, castrated 3/20/61, at 139 days of age.

reached maximum during the test given eleven days following the final
androgen injection. A rapid decline then set in for all measures, first
mounting, then abortive mounting, followed by nudging and nosing, in that
order. Pre-treatment levels of performance reappeared some 64 days after
the final injection of testosterone.

(3). Ram # 13.

In the earlier discussion of the development of sexual behavior, the history
of the attenuated sexuality evidenced by # 13, castrated 20 March, 1061 at
the age of 118 days, was presented. Tt was obviously of interest to determine
whether androgen treatment would have an effect upon the behavior of
this ram.
It can be seen from Fig. 7 that the only measure recorded for # 13
between February and August, 1962 was a low frequency of nosing activity.
The pre-treatment series begun in October, 1962 continued to reveal an
absence of the major parameters of sexuality characteristic of this species.
At weekly intervals, beginning r5 October, # 13 was injected with 100 mg.
of testosterone, IM. In contrast to the injection schedule of the two previ-
ously discussed rams, #13 was injected once per week for five consecutive
weeks.
274

An abrupt increase in nosing occurred after the first injection. However,

it was not until after the third injection that nudging appeared during tests
of # 13. Mounting was observed once, two days after the fourth injection,
and twice, two days following the fifth and final injection. The frequency
of abortive mounting mimicked that of mounting. Both of the latter measures
declined to zero 16 days, and nudging dropped out 30 days after the final
injection.

Fig. 7. Record of ram #13, castrated 3/20/6i, at m8 days of age.

During test sessions occurring when # 13 was receiving exogenous an-

drogen, he evinced considerably more responsiveness to the estrous ewe
than he had previously. However, his courtship activity resembled that of
other rams very slightly. Prominent in the motor activity of # 13 was the
intention bunting that ended in a nudge described previously in the presenta-
tion of the data for # 21. When mounting occurred, penile erection was just
barely visible to the observer, a result, doubtless, of the long absence of
hormonal support for the maturation and maintenance of the penis.

(D). DISCUSSION OF ANDROGEN REPLACEMENT THERAPY

Within the narrow limits of this experiment, several comments are pos-
sible. Androgen treatment resulted in a substantial increase in the frequency
of display of components of sexual behavior in each of the three rams.
 275

The form of testosterone used was one having long-acting properties, when
used in replacement therapy in man. The dosage of ioo mg. per week was
necessarily arrived at arbitrarily, there being a surprising lack of information
regarding supportive androgen therapy in this species, as contrasted with
information available from other forms used in this type of experiment.
In the treatment of eunuchism in man, the suggested dose of Depo-Testo-
sterone is 200 to 400 mg. injected at intervals of three to four weeks. The
animals varied in weight during the treatment period, e.g. # 21 - 122 lbs,
# 18 - 142 lbs, and # 13 - 117 117S.It is doubtful that differences in response
were the result of a weight-dosage factor, although a more extensive series
of animals and dosages would be needed to clarify this point.
The record of # 21, a sexually experienced adult, following castration
revealed an orderly sequence of disappearance of adult sexual components,
e.g. copulation (mounting with intromission and ejaculation), nudging, and
some decline in total nosing frequency, in that order. Responses by # 21
to an estrous ewe in the test situation were essentially nonexistent within
52 days after castration. By way of comparison, in the maximally ex-
perienced male cat, post-castrational decline of sexual behavior varied from
an almost immediate disappearance of intromission in one individual, to
intromissions performed three and a half years after castration in another
(ROSEN1LATT & ARONSON, 1958). It is now well established that vari-
ability in behavioral response to castration as a function of individual
genetic, experiential, and hormonal factors exists in all vertebrate species
examined.
The unfolding of the sexual repertoire under stimulation of exogenous
androgen in #21 occurred beginning with increases in frequency of the
appetitive components, nosing and nudging, and then mounting. Intromission
was observed, but behavioral evidence of ejaculation was not recorded. The
maximum frequency of mounting with intromission (15/10 minutes) was
very much in excess of maximum scores provided by normal males tested
under the same circumstances (8,po minutes). It may well be that such
hyperactivity reflects an overdosage of the injected hormone. It is clear that
the frequency of abortive mounting which rose to a maximum of 12/10
minute test can be attributed to the generally high level of excitement of
#21 during test sessions. Decline in all measures to pre-treatment levels
was complete about two months after the final injection. Rather than a
gradual phasing out of each component, waning of appetitive and consum-
matory acts occurred simultaneously.
The results obtained from # 18 differed from those of # 21 in several
respects. Frequencies of nosing and nudging rose more precipitously in # 18
276

and measures of mounting and abortive mounting declined to pre-injection

levels prior to those of nudging and nosing. The reasons for these dif-
ferences, assuming they are of some significance, are not clear. Both had
had successful experience in mating prior to the treatment, # 21 more so
than # 18. Both had been castrated, # 18 for a much longer period of time
than # 21, and both had received the same amount of injected androgen. It
might be suggested that these results represent an example of individual
variation in sensitivity to the exogenous hormone, but the available data
do not allow for a rigid stand to be taken on this point.
Of the three rams used in the experiment, # 13 provided results of most
interest. This ram, castrated at 118 days of age, had never displayed sexual
behavior in tests administered over a period of one and one half years. His
general behavior in the flock situation was one of extreme submissiveness
and further, # 13 had never been observed to mount another male during
play activities. It may be recalled that this latter feature was one of the
common social patterns displayed by all other young rams in the unisexual
flock. The injection of androgen evoked a marked increase in the frequency
with which #13 attained physical contact with stimulus females, as reflected
in the rise of the nosing score. Nudging was manifested after the third
injection and mounting appeared after the fourth, at which time a very sub-
stantial increase occurred in nudging frequency. The virtual absence of the
later components of sexual behavior prompted the administration of a fifth
injection of androgen to # 13. A response of two mounts?i0-minute test
was obtained. Upon withdrawal of hormone support, mounts, abortive
mounts, and nudging declined to zero, but nosing frequency remained at
levels somewhat above those manifested during pre-injection tests.
The data thus indicate that the full gamut of sexual behavior, of which this
ram was capable (all, that is, except for intromission and ejaculation) were
evoked by the hormone treatment. As noted previously, his behavior was
qualitatively distinct from that of the other two rams. The precision of
movement with which a normal ram courts the ewe was not evidenced. In
quantitative terms, # 13 could not be said to have attained levels of per-
formance approaching those of normal males. Arousal of sexual interest
accompanied hormone support, but the concomitant motor activity was quite
ambivalent.

SUMMARY
1. A generalized description of the motor components comprising male courtship is
presented. Included among the major acts are a) nosing the perineum; b) nudging, a
complex of several subacts including orientation behind the ewe, extension and flexion
of one foreleg, tilting and lowering the head accompanied by a low-pitched vocaliza-
 277

tion and flicking of the tongue ; c) the Flehmen or lip curl, as displayed by the ram
is described, but the signal function in this species is problematical; d) mounting ac-
companied by e) pelvic oscillations, f) intromission, and g) ejaculation, defined beha-
viorally as an especially deep thrust After-reactions may take the form of remounting
the ewe, leaving to court another female, or standing while the just-mounted ewe leaves.
2. The motor components displayed by the ewe vary with its hormonal condition.
When anestrus, the female manifests either passive or active avoidance depending upon
the persistence of the ram. During the 15 to 28 hours of estrus, female motor activity
ranges through a spectrum of low to high to low intensity responses. For the first 3-5
hours, low intensity acts include standing, head lowered and pinnae of ears somewhat
flattened, swinging the head back to watch the courting ram, walking off and then
standing and looking back at the ram. Persistent nudging by the ram at this time
results in acceptance. From about the 5th to the 15th hours of estrus, the ewe is notably
more aggressive. Soliciting behavior takes the form of approaching the ram, nuzzling
and pushing its head into the flank and scrotal regions. Low intensity responses super-
vene from the 15th hour to the end of estrus, grading into the avoidance behavior
characteristic of the anestrous ewe.
3. Observations of a small, closed flock comprised of 6 ewes and 1 ram in an indoor
pen provided information on dominance relationships among the ewes and the influence
of rank upon female sexual activity. Additional information obtained from these obser-
vations included evidence of memory by the ram of the identity of females with which
he had mated.
4. The ontogenesis of male courtship acts was studied in a flock of 8 rams, 4 of
which were unaltered, 2 castrated (one at 118 days, the other at 139 days of age), and
2 deprived of heterosexual contact until well past the age of physiological maturity.
All unaltered rams achieved copulation, but only after from 10 to 20 ten-minute expo-
sures to a stimulus ewe. One of the castrate rams succeeded in mating after 12 expo-
sures, the other displayed no courtship behavior at the 20th exposure at which time he
was 540 days old. Of the 2 female-deprived rams, one achieved a mating at the first
exposure to an estrous ewe; he was 419 days old at the time. The other deprived ram
evidenced successful courtship on the 12th exposure when he was 676 days of age.
General observations of the ram flock revealed that all components of the male pattern,
except for the consummatory acts of intromission and ejaculation were manifested
during social interactions in the flock. Such behavior was displayed during the play
activities of all except one of the castrate rams.
5. Three of these rams were further examined with the view of establishing the
effect of injections of male hormone. Included in this study were the two castrates and
one unaltered ram from the original flock of eight. The latter was castrated after
having developed consistent, adult sexual behavior. Treatment began after all three
rams displayed a decline to little or no sexual behavior in a test situation. In all three
instances, treatment with exogenous androgen evoked the complete male courtship pat-
tern, although one of the original castrates appeared to be almost refractory to the
treatment.
6. A discussion of these findings is included in each of the various sections of this
report.

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- (1956). Characteristics of masculine "sex drive". in: Nebraska Symposium on Mo-

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GEIST,V. (1963). On the behaviour of the North American Moose (Alces alces ander-
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ZUSAMMENFASSUNG
1. Ein allgemeiner Überblick über die motorischen Komponenten der Werbung des
Schafbockes wird vorgelegt. Die Werbung besteht aus folgenden Hauptakten : a) Be-
schnüffeln des Dammes; b) Aufforderndem Anstossen (nudging), einem Komplex ver-
schiedener Unterhandlungen wie Orientierung hinter dem Schaf, Ausstrecken und
Beugen eines Vorderlaufes, Neigen und Senken des Kopfes, begleitet von einer tiefen
Lautäusserung und Hin- und Herbewegen der Zunge; c) dem Flehmen des Bockes,
dessen Signalfunktion bei dieser Art noch problematisch ist; d) dem Bespringen, be-
gleitet von e) Beckenoscillationen, f) Intromission und g) der Ejakulation, die ver-
haltensmässig als besonders tiefer Stoss definiert ist. Nachreaktionen können in wieder-
holtem Bespringen des Schafes bestehen, Umwerben eines anderen Weibchens oder
Verweilen, während das soeben gedeckte Schaf sich entfernt.