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A Study of Some Factors
Influencing Fertility and Sterility

in the Bull

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BY

HERBERT L. OILMAN

Reprinted from the
Annual Report of the New York State Veterinary College

1921-22

... • • ». ' • •

A Study of Some Factors
Influencing Fertility and Sterility

in the Bull

A THESIS

Presented to the Faculty of the Graduate School

of Cornell University for the Degree of

Doctor of Philosophy

BY

HERBERT LESTER OILMAN, D. V. M., M. S.

ii

Reprinted from the
Annual Report of the New York State Veterinary College

ALBANY

J. B. LYON COMPANY, PRINTERS

1922

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'1

A STUDY OF SOME FACTORS INFLUENCING FERTILITY
AND STERILITY IN THE BULL

HERBERT L. OILMAN"
Veterinary Experiment Station, Cornell University

Normal reproduction is the fundamental foundation upon which
the entire cattle industry rests. For this reason, any factor
capable of interfering with it is a detriment to the industiy, and
a matter of prime importance to the breeder and the veterinarian.
With the relative increase in number and value of cattle, an.l the
fact that the profession is depending more and more on this in-
dustry for a livelihood, these problems are assuming greatei- im-
portance. The part plaj^ed by the bull has been emphasized
entirely too little, with the result that, as in human medicine,
many fail to appreciate the effects of sterility or lowered fertility
in the male. The part played by the sire in the spread of genital
infections, though discussed frequently, has received little sys-
tematic investigation.

The bull miLst be regarded as at least half the herd, not only
from the standpoint of the characters he imprints upon his prog-
eny, but because of his relation to the reproductive efficiency in
the herd. It seems quite probable that he does disseminate during
copulation, infection associated \\i\h. the genital organs, with tlie
result that the bull is a very important factor in a study of the
subject. Too frequently, his ability to copulate in an apparently
normal manner, is taken as a standard of fertility. Gross changes
in his genitalia, or the absence of spermatozoa from the semen
are given due consideration, while other more ol^seure al)nor-
malities are not looked for nor regarded in their proper light.
Neither fertility nor sterility are always absolute, but the terms
should be used relatively inasmuch as we may have all degrees of
infertility or impotency. All Too frequently we foi'get the many
delicate and intricate mechanisms involved in ihe reproductive
process, with the result that many phases of the problem are neg-
lected or disregarded. The genital organs woi'k as a unit, each
part of which must function in perfect accord with the others to
the end that full fertility may result. The physiological factors
involved in the formation of the semen are too little understood,
or at best, our knowledge regarding them is more or less hazy.

The purposes of the present work have been: (1) to summarize
the work so far done on the subject. (2) to review brietly the
known facts throwing light on the anatomy and physiology of the
male genital organs, (3) to carry out systematic studies upon the
pathology and bacteriology of the genital tract of the bull, and
(4) to ascertain if possible whether the ])ull is a disseminator of
those infection!^- which interfere with reproduction in the female.

f3l

51>7778

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Tlie work has been carried on for the most part from the point
of view of a laboratory man cooperating with clinicians. No
attempt is made in this paper to give detailed clinical data, methods
for physical examinations, etc. There are included many state-
ments "and some data given in a preliminary article on the sub-
ject. While the subject is broad in its scope, in fact too broad for
great detail, it is hoped that a start has been made toward future
and more detailed investigations.

History
References to, and investigations relating to, the part played by
the bull in the process of reproduction in the herd, and in the
spread of genital infections, have been limited largely to those
phenomena caused by Bad. ahortum. Bang (1) originally called
attention to the possibility of the male transmitting the organism
discovered by him, but he reached no definite conclusion on the
subject. James Law (2) writing on contagious abortion in cows,
early suspected this possibility when stating under " casual infec-
tions," that — " In a case which came under the observation of the
Avriter recently, a family cow, kept in a barn where no abortion
had previously occurred, was taken for service to a bull in a herd
w^here abortion was prevailing, and though she Avas only present
at the latter place for a few minutes, she aborted in the sixth
month." Jansen, as quoted by Sand, reports the case of a cow
from an aborting herd having been taken into a herd that had
been previously quite free from the disease. Soon after her arrival
she aborted, and later cow after cow of the original herd aborted.
The owner kept the matter a secret, and sent his cow to a neigh-
bor's bull for service, with the result that for two years abortion
prevailed among cows served by this bull. McFadyean and Stock-
man (3) later, in experimental work, attempted but failed to in-
fect cows by using a soiled bull for service. Hadley and Lothe
(4) state: "A large number of stockmen hold that the bull is an
important factor in the transmission of contagious abortion in
herds. A smaller number believe that the bull merely acts as a
passive carrier of the abortion disease and is not actively concerned
in the transmission." In a subsequent bulletin, Hadley (5) re-
marks : ' ' The abortion organisms may enter the body

during sexual intercourse." In an experiment carried on by the
same author and co-workers, abortion-free virgin heifers were mated
to abortion-infected bulls, infection being evidenced by positive
reactions to the complement fixation and agglutination tests. His
results indicate, he believes, " that the bull is not so important a
factor in transmitting abortion as many believe." The con-
clusions are: "Bulls may become infected with abortion bacilli.
Bulls that reacted to the blood tests were incapable of dissemi-
nating the abortion disease to the abortion-free heifers with which
they were mated. Bulls appear to possess a sexual or individual
immunitv to abortion infection that renders them less susceptible

than cows and induces a milder form of the disease. The resist-
ance appears to be due to certain anatomic and physiologic differ-
ences in their Sexual organs which make them less favorable places
for the growth of the abortion germs than those of the opposite

sex."

Buck, Creech, and Ladson (6) applied the agglutination test to
325 mature bulls, of which 288 were negative and 37 positive.
Bacillus abortus Avas isolated from five animals, of which three
showed marked lesions, two in the seminal vesicles, and one in the
left testicle. They conclude: " B. abortus may involve organs of
the generative apparatus of bulls, producing chronic inflammatory
changes. Of the generative organs, the seminal vesicles appear to
furnish the most favorable site for the lodgement and propagation
of abortion infection."

Schroeder and Cotton (7) cite the case of a bull which reacted
to the abortion test and, on post mortem, Bact. dbortum was
isolated from an abscess of one epididymis. They state: "Our
attempts to produce a similar case of infection artificially failed,
and, in agreement with the difficulties many investigators have
had to obtain incriminating CAddence against bulls, we have thus
far failed to infect bulls in any way that justifies the assumption
that they are important factors in the dissemination of abortion
disease." Further, they conclude: " Regarding the dissemination
of abortion disease by bulls, we may say, however, that it would
be foolhardy in the dim light of our present knowledge to take
liberties with reacting bulls, or bulls from infected herds, or
promiscuously used bulls."

Cotton (8)"^ failed to demonstrate the presence of abortion bacilli
in the genital organs of the bull used to serve aborting cows, or in
the testicles of two bull calves, one of which had been fed and the
other injected M'ith the cultures of the abortion bacillus. He con-
cludes that the bull does not harbor the organisms in the testicles.
Carpenter (9) injected both streptococci and Bact. abort um into
the scrotal sacs of young calves, and intravenously in others. In
no case was he able to recover the organisms from any part of the
genital canal, except for a streptococcus in one instance. Tiettger
and AYhite (10) were unable to obtain evidence of the presence of
Bact. abortum in three bulls slaughtered after repeated reactions
to the complement fixation and agglutination tests. The three
bulls had been under observation for three years, with no conclu-
sive evidence to indicate that they were a source of danger to the
herds in which they were a part. They believe that the bull trans-
mits the infection as a passive carrier.

Attempts at artificial inoculation by natural channels have
failed, with the possible exception of McFadyean, Sheather, and
]\rinett (11) who were able to infect the bull by the prepuce in two
cases and by the mouth in one case. The results, however, are by
no means conclusive. They conclude, nevertheless, that catth; of
any age of either sex may be infected by natural channels with the
bacillus of epizootic abortion.

Schroeder (12) carried out investigations to ascertain the fre-
quency \\'ith wliicli bulls react to abortion tests, and the frequency
Avith which lesions chargeable to al)ortion bacilli occur in the re-
productive organs of reacting bulls. Studies were also pursued
which he states conclusively prove that bulls with infected repro-
ductive organs may expel abortion bacilli with the seminal fluid.
In the first two mentioned investigations 325 bulls from a Wash-
ington abbatoir were tested, and slaughtered upon reaction. "Ap-
proximately ten per cent of the bulls reacted, and approxi-
mately ten per cent of the reacting bulls showed lesions of the
reproductive organs from which abortion bacilli were isolated.'*
The value of these studies, he emphasizes, lies not in " that they
give us a measure of the proportion of bulls that react positively
to abortion tests or the proportion of reacting bulls that are car-
riers of abortion bacilli," but in " the fact that they show that
abortion bacillus disease of tlie bull's reproductive organs is not a
whoU}' unique affection which practically may be ignored, but an
important condition that must be taken into account in our efforts
to combat infectious abortion, since it has been proved to be asso-
ciated with contamination of the seminal fluid." In discussing
the method by which infected bulls transmit the organisms to
cattle, he believes that leakage of semen from the penis, or vaginas of
cattle after service, contaminates the food which subsequently gains
entrance to their digestive tracts. As the result of a series of

experiments, he states: " the results fail to justify in the

least degree the assumption that cows are infected with abortion
bacilli via their vaginas or uteruses at the time of copulation, or
that the bull, through copulation, is an agent in the spread of
abortion disease."

The work so far alluded to, has been limited to infection with,
and the transmission of, Bart, ahortnm and the lesions associated
with such infection. The last mentioned author, however, states :
"A search for other specific causes of al)ortions among cattle
has not been neglected, and bureau investigators could relate at
great length stories similar to those which investigators have told
about microorganisms isolated from the products of abortions and
the uteruses of cows that have aborted. Bacilli of various kinds,
different types of micrococci, and spirilla or vibrio have been found
repeatedly ; but when their pathogenicity has been tested in accord-
ance with widely recognized and accepted and required bacterio-
logical standards, not one shred of evidence has been obtained to
prove them true etiological factors of bovine abortions. What role
such microorganisms may have as causes of the sequellae of in-
fectious abortions, and of other, possibly, independent, abnormal
processes in the reproductive organs, is far from clear and merits
careful study." Iladley (5) mentions the fact that: " Unques-
tiona1)]y the male often ])ecomes infected with the germs that
produce the various secondary' diseases in the female, Avhich are
properly classed under the more inclusive term 'abortion dis-
ease.' " Also, speaking of the rarity with which the bull acquires

abortion infection, he alludes to the i'aet that he may act as a
" meclianical carrier of various disease germs from an infected to
a healthy cow." Carpenter (9) working on the female genital tract,
comes to the conclusion that, in all probability, the genital organs
are normally free from bacteria. Barney (13) quoting Huet finds
that bacteria may be present in the seminal vesicles of healthy
animals (horses, cattle, pigs, and laboratory animals). This, he
states, corresponds with the well recognized ilndings in other parts
of the genito-urinary tract, not only in animals, but in man. He
(Huet) has further found that in animals dying of acute septi-
cemia, the specific organism (anthrax, i)neumococcus) is to be
found in the vesicular secretions. Furthermore it was definitely
shown that an infection could be transmitted to the female during
the act of copulation.

Williams, AV. L. (14), calls attention to the lack of vet-
erinary literature relating to the pathology and bacteriology
of the male genital tract, except as related to infection with
Bad. ahortum. Infection with other types of bacteria is empha-
sized, the clinical recognition of such, with the accompanying
pathological changes, and of the numerous phenomena involved in
tlie process of reproduction in the male. The semen and its essen-
tial germinal elements are taken up with reference to the entire
lack of study devoted to them, and some of the abnormal changes
are described. In a later contribution (15), he takes up the part
played by the bull in the dissemination of genital infections and
.states: " Clinical studies now indicate Avitli great clearness that
the bull is an active spreader of that group of genital infections
which cause sterility, abortion, and related phenomena."

Williams, W. W. (16) studied the semen with reference to steril-
ity, emphasizing the importance af its examination in the diag-
nosis, giving methods for collecting samples, staining' of sperms,
and some of the abnormalities encountered. The work is funda-
mental, and shouhl be of great practical impoi-taiice to all interested
in the proljlem. In a later paper (17), he brings out a more ex-
tended discussion of the question. He concludes that the clinical
examination is of vital importance. v.r\(\ that the efficiency of the
semen depends not only upon its physical properties Init upon thc-
number of spermatozoa that are motile, the degree of motility,
degree of obligospermia, and the percentage of imperfect spermato-
zoa, either deformed or immature. Of forty bulls examined, he
finds that twenty, or fifty per cent, sliowed lessened fertility, and
others, aside from this, showed minor clianges in the genital
organs or semen. The same author suliseciuently takes up the sub-
ject of reproduction from the viewpoint of l)oth sexes, but emplui-
sizing infection in the male, and the frequency with which lowered
A'itality of the germinal cells occurs. Hopper (18) states: "A
diseased bull may manifest non-fertility or decreased potency in
diiTerent ways — by repeated service to apparently normal females
without conception, by a high a])ortion rate in females that have
been apparently normal, by characteristic infections following! the

use of any particular sire, or by abnormalities in the breeding
tract noted by rectal or physical palpation."

The observations of Williams (19) in a pure bred dairy herd
bring out quite clearly the relation of the bull to the dissemination
of genital infections. The bulls in this particular herd were ab-
normal in many respects, as demonstrated by pathological changes
in their genital organs, bacterial invasion of the parts, abnormali-
ties of the semen and spermatozoa, and the probable transmission
of infection to the females. Several of the sires from this herd
furnished much of the material for the early basic work of this
investigation. Since then the tracts of other sires have been worked
upon with quite similar or identical results.

To summarize the work already done, most investigators have
considered the bull as merely a mechanical carrier of Bact. abor-
tum infection, though all are more or less suspicious of his ability
to become an active spreader. Schroeder, however, states that the
organisms are eliminated Avith the semen, but infection of the fe-
male occurs secondarily through the digestive tract by contamina-
tion of the food with the semen. Other investigators bring out
fundamental points demonstrating the importance of other organ-
isms than the Bang bacillus and call attention to the need of a
more thorough study of the anatomy, physiology, and pathology
of t.he male genital tract.

Any study of the genital organs must of necessity rest funda-
mentally upon a thorough knowledge of the anatomy and phy-
siology of those parts. Too few of us have stopped to consider
thei^e questions thoroughly, with the result that our ideas on the
problem are more or less vague. It is much easier to understand
why abnormal spermatozoa occur so frequently, or changes take
place in the semen with death or weakening of the germinal ele-
ments, if we realize or stop to consider the highly differential pro-
cess of spermatogenesis, and the various structures which contrib-
ute to the formation of the semen. AVe must come to realize that
each part of the genital tract is essential to the normal functioning
of the whole, and that the genital tract and rer)roduction are in
turn dependent upon the proper functioning of the entire body.

Walker (20) emphasizes the importance of a thorough knowledge
of physiology' in stating, ''Although the subject of sterility has
attracted the attention of the medical profession; and although
much has been written on its causes and treatment, it cannot be
claimed that the practical results olitaincd up to the present time
are satisfactory, or that when consulted for sterility, the medical
man of today can hold out to his patient much more hope of suc-
cessful treatment than the medical man of fifty years ago. Our
failure in this respect is in the main due to an ignorance of the
physiology of reproduction."

To bring out some of these points, the anatomy of the tract will
l)e reviewed briefly, together with the physiology of reproduction,
and the various factors which should be considered in a studv of
the problem.

Anatomy and Physiology

In origin and early development the ovary and testis are identi-
cal. The gonad and mesonephros or primitive kidney are de-
veloped from the urogenital fold. The gonad first forms as a
medio-ventral thickening of the fold, which gradually expands
until it becomes attached by a mere stalk. At first, the gland is
made up merely of a superficial epithelial layer, and an inner
epithelial mass, or epithelial nucleus. In the process of develop-
ment, large primordial germ cells migrate from the entoderm of
the future intestinal canal, and pass through the stalk to the gonad.
In the case of the male gonad, seminiferous tubules are very
difficult to make out in eml)ryos smaller than 24 millimeters. Then
they suddenly differentiate out as solid cords of germ cells, -while
the connective tissue grows in around them. These connective
tissue sheaths unite at the center of the organ to form the anlage
of the mediastinum testis. The testicular tubules unite and con-
verge toward the hilus, there to meet the anlage of the rete. At
the mesonephric end of the testis, the rete first appears as a col-
lection of cells, differentiating out from the inner epithelial mass
of the gonad. These cells gradually grow out to meet the collect-
ing portions of the mesonephric tubules on the one hand, and the
seminiferous tubules on the other. The rete is represented as
cords of cells at first, which in forty millimeter embryos hollow out
to form tubules.

The mesonephros, or primitive kidney, ea;'ly starts to degenerate
cranio-caudally, — the tubules becoming separated into a cranial
and caudal group. The collecting and secretory parts of the
cranial group separate, the collecting tubules growing out to meet
the rete with wliieh they unite to form the efferent ductules of the
epididymis. The caudal group of tubules is vestigial and becomes
the paradidymis. The mesonephric duct becomes the vas deferens,
connecting as it does with the tubules of the epididymis, and
emptying into the urethra at Mliller's tubercle or, as it later be-
comes, the colliculus seminalis.

The seminal vesicles arise as hollow saccules from the dorsal
wall of the mesonephric duct just as it empties into the urethra.
The prostate develops aa an outgrowth of the dorsal urethra .just
posterior to ]\[iiller's tubercle. The bulbo-urethral glands appear
as solid, paired, epithelial outgrowths from the entoderm of the
urogenital sinus.

!Miiller's duct, at first a solid tube growing from the anterior
part of the mesonepliros, and ending at ^Midler's tubercle, becomes
a hollow tube, and in the female forms the entire genital tract ex-
cept for the gonad and the lower part of the vagina. In the male,
the anterior part remains as the vestigial appendix testis, and the
posterior part, as the vagina masculina, Ellenberger states, how-
ever, that this embryonic structure is very seldom seen in the
mature bull.

10

The Male Reproductive Organs include the penis and testes,
together with the excretory passages which connect the testes with
the urethral canal. These excretory duets include the epididymis,
vas deferens, and seminal vesicles. Posterior to their termination
in the urethra, there aro connected the ducts of the prostate gland
and the bulbo-urethral or Cowper's glands.

Testes : The testicles of the bull are relatively large. Vaiying
with the size and age of the animal they measure from fourteen to
seventeen centimeters in length, including the epididymis, and
from six to eight centimeters in diameter. Each testicle is en-
closed within a serous sac, the tunica vaginalis, whose
visceral layer is very intimately fused with the underly-
ing covering of the organ, the tunica albuginea. The tunica
albuginea is quite thin and consists of connective tissue
which is rich in elastic fibres. Muscular tissue is not present as it
is in the case of many mammals. Inside the tunica, and closely
attached to, though separated from, the parenchyma by a thin
layer of connective tissue, is a laj-er of very loose connective
tissue, M'hich because of its rich supply of blood vessels is termed
the tunica vasculosa. The parenchyma is of a yellowish gray
color, and of a rather soft consistency. It is made up of the semi-
niferous tubules, rete, and the connective tissue stroma, the medi-
astinum testis. On section, the mediastinum appears as the center
or axis of the entire organ. It is star-shaped, and radiates connect-
ive tissue septa out into the parenchyma to support and separate the
tubule.^. Ellenberger states that the testis of the bull and all
ruminants lacks a closed system of interlobular septa, because of
the feeble development of the connective tissue.

The principal blood vessels and rete tubules are found in this
structure, the function of the latter being to connect the seminifer-
ous tubules and the efferent tubules of the epididymis. The
epithelium of the rete is quite irregular. — consisting in places of
a single layer ; in others, of two layers. At some points there are
formed groups of several cells lying over one another, with swollen
liomogeneous basal cells, which sometimes form projections into the
lumen.

The interstitial tissue, besides conveying the blood vessels to the
organ, contains many " interstitial cells.'-' These cells are rela-
tively sparse in the adult bull, and are comparatively delicate,
slightly granular, often shuttle-shaped, with a rather small nucleus.
Embryologicnlly they are derived from a syncitium arising from
the mesothelium of the genital ridge, differentiating out by growth
of the cytoplasm. They contain large quantities of fat, and
elaborate the internal secretion of the testis. This secretion gov-
erns the development of the secondary sexual characters, and has
a profound influence on the general body metabolism, and develop-
ment of the skeleton. The interstitial cells appear early in
embryonic life even before there is any differentiation of sex, and

11

their greater relative developmeut iu the fetus is indicative of a
future male development. In very young embryos, the growth is
very rapid, followed, however, by a period of atrophy, during
which the seminiferous tubules undergo marked development.
Pende (21) states: " There seems to be an inverse relation be-
tween the growth of the tubular and interstitial tissues, as one is
hypoplastic when the other is in full activity." From birth to the
onset of sexual maturity, whicli may be called a period of rest for
the testicle, the cells are few in number. With the accentuation of
the secondary sexual characters, and the beginning of sexual life,
these cells again increase in number and acti^-ity.

The parenchyma of the testis consists for the most part of the
seminiferous tubules, which, on account of the courses they take
in the different regions, are divided into groups. The peripheral
tubules are the much-contorted tubuli contorti. These anastomose
to form the mucli shorter tul)idi recti. These in turn anastomose
frequently, uniting to form the rete testis. The rete proceeds
through the mediastinum to form the efferent ductules which
break through the tunica albuginea to form the greater part of the
liead of the epididymis. The tubuli contorti are the longer and
more numerous of the tubules, for it is liere that practically all
the spermatozoa are produced. The straight tubules are relatively
so short that they may be regarded more in the light of the begin-
ning of the system of excretory ducts.

The seminiferous tubules consist of a thin peripheral meml)rana
l)ropria upon which rests tlie seminal e])ithelium, which is made up
of the essential semen forming cells, and the cells of Sertoli. The
spermatogenic cells may be divided into three groups, from within
outward: the peri])heral single layer of small cuboidal spermato-
gonia ; one or two rows of large spermatocytes ; and three to five roAvs
of spheroidal spermatids. Tlie cells of Sertoli are more or less of the
syncitial type. — large in size and irregular in outline. They occur
at various intervals between the layers of spermatogenic cells, with
their bases resting upon the mcmbrana proi)ria. Centrally they
send out protoplasmic processes for variable distances,— some
even reaching the l)order of the innermost cell layers.

Spermatogenesis: In this process, the ])rimary germinal cells,
the spermatogonia, divide to form the i)riniary spermatocytes,
^laturation consists of two cell divisions of tlie primary sperma-
tocytes, and these in turn form four spermatids. During the pro-
cess, the number of chromosomes is reduced to half the number
characteristic of the species. The spermatids then become con-
verted into mature spermatozoa. Tliis mode of transformation
may be seen in Plate I. In the process, the nucleus of the sperma-
tid forms a large part of the head; the eentrosome divides, part
passing to the extremities of the neck. One eentrosome ])ecoraes
the .interior, and remains altaelied to the head, while tlie other
divides to form the posterior eentrosome. The latter is divided
into the anterior part, and the posterior nodule or annular ring.

12

Besides this, the posterior eentrosome becomes elongated to form
the axial filament, and the cytoplasm forms the sheaths of the neck
and tail. The spiral fihiment of the connecting piece is derived
from the cytoplasmic mitochondria. At this time, a large part of
the cellular cytoplasm is cast off. Meanwhile, the spermatozoa
sink their heads into the long protoplasmic processes of the Sertoli
or " nurse " cells which furnish nutritive material for their com-
plete development. Finally the adult cells are cast off into the
lumen. The structure of the spermatozoa, and a discussion of the
semen will be taken up later.

Epididymis: The epididymis is divided into three parts: the
head, body and tail. The head is made up principally of the
lobules formed by the much-coiled efferent ductules proceeding
from the rete. The ductules, about twelve in number in the bull,
unite to form the body, which remains coiled and runs along the
postero-medial part of the testicle to which it is more or less closely
attached. To quote Ellenberger (23) : " The transition from the
rete into the ductules is gradual, as the characteristic epithelium
of the latter (ductules) begins in cavities without walls, and at
first, gradually form a wall which is well marked out as a thin

ring of interstitial tissue The epithelium of the ductules is

in sharp contrast to the rete in that it has a single-layered ciliated
columnar epithelium, in which here and there one finds round basal
cells. The dark and light columnar cells alternate; the cilia are
often cemented together, and form cone-shaped, homogeneous
appearing protuberances. The secretorj^ activit}^ is quite clearly
observable. In the light cells one finds secretory globules, accumu-
lating in rows, sometimes above, other times below, arches of cells.
The secretory droplets pass from the cells into the lumen, and
often lie in irregular layers on the epithelium ; also the basal cells
appear swollen and shoved out between the cylindrical cells." At
the lower extremity of the testicle, the tail is formed, which is
globular in shape, and more or less loosely attached to the testicle.
Here the ductules anastomose freely, gradually become less coiled,
and end in the single excretory tube, the vas deferens. The epithe-
lium at the tail part is more or less of the pseudo-stratified
columnar, ciliated variety. Outside this is a membrana propria,
a circular muscular layer, and a connective tissue coat. The secre-
tory activity is very marked here and one finds much secretion in
the lumen. Courrier (24), working on the bat, suggests that the
glandular activity is conditioned by the secretion from the inter-
stitial (endocrine) gland. The action of the secretion is to dilute
the large mass of spermatozoa present, nourish them to some extent
and also stimulate them to active motility. Stigler (25) states
that the properties of the sperms are modified in the epididymis;
the motility, the ability to resist heat, and other properties are
augmented, at least in the case of the guinea pig, rat, and mouse.
Some authors state that the sperms first become motile when in
contact with the prostatic secretion, but I have repeatedly exam-

13

ined the contents of the tail of the epididymis of the bull, rabbit,
and guinea pig, finding full motility in each case, though the dura-
tion is not nearly as long as when the sperms are ejaculated in the
semen.

The Vas Deferens is quite narrow (2 mm.) and runs from the
tail of the epididymis to the verumontanum, or colliculus semin-
alis, w'here it empties into the urethra in common with the duct of
the vesicle. At first it is lined by epithelium similar to that of the
vas epididymis, but this changes over into a peculiar low stratified
type. Ellenberger describes it as follows: " The epithelium
shows a very pronounced basal coat. The overlying cell zone
shows more (at the most, three) rows lying over each other of
elongated nuclei, while an outline of cell form is not ordinarily
noticeable, so that it may be spoken of as a syncitium, and at the
same time as a many layered epithelium." The mucosa forms low,
broad folds into the lumen. The tunica propria is a thin con-
nective tissue layer. The submucosa consists of thin connective
tissue. Three muscular coats are present : an inner thin longitud-
inal layer, middle circular layer, and an outer longitudinal layer.
All are more or less intimately blended, and are surrounded by
the adventitia, made up of connective tissue, elastic fibres and
scattered longitudinal muscle cells of the internal cremaster mus-
cle. Near the dorsal siu*face of the bladder, the ducts come in
close apposition, and for ten to twelve centimeters dilate to form
the ampullae. Here the mucous membrane bcomes much plicated,
forming long folds which anastomose freely. The function of the
vas is to convey the spermatozoa and secretions from the epididymis
to the urethra. Disselhorst (26) believes the ampulla acts "as a
seminal reservoir and states that he has found spermatozoa stored
up in the little pockets in the walls of this structure in animals
during the rutting time. He suggests, further, that there is a
relation between the state of development of the ampulla and the
time occupied by copulation. When the organ is small or absent,
as in dogs, cats, and boars, the coition is a slow process, but when
the ampulla is large and well developed, as in horses and sheep,
the coitus requires a relatively short time. Inasmuch as coitus is
so rapid in the bull, and the ampulla is so well developed, it seems
as though this function is very probable.

The Seminal Vesicles are very compact glandular structures
^ylng on either side of the median line, on the dorsal side of the
'ladder, and ventral to the rectum. In the mature bull they
rlieasure ten to twelve centimeters in length, four centimeters in
width, and about two and one-half to three centimeters in thick-
ness. The glands are distinctly lobulated. quite tortuous, and are
often asymmetrical in size and shape. They converge posteriorly,
to empty into the urethra at the colliculus seminalis with the
ampulla, in a slightly oval slit in the mucosa. IMicroscopically,
the gland is of the anastomosing tubular type, with very poorly

14

developed excretory ducts to the glandular cavities. Posteriorly
one finds centrally a few sinus-like narrow excretory passages,
which open into the somewhat larger collecting and excretory duct.
The epithelium is of the simple columnar tj-pe and produces a
relatively large amount ol' secretion. The gland cavities are sur-
rounded by a membrana propria, over which is a relatively thick
layer of smooth muscle. Outside this is a connective tissue covering
which sends traljeculae or septa in between the lobules. The secre-
tion of the seminal vesicles is a tenacious albuminous tliiid with
a slightly yellowish tinge, all or part of which appears in the
ejaculate in the form of swollen sago-like grains which are soon
dissolved following ejaculation and the liquefaction of the semen.
The proteid compounds belong to the group of histones. The
secretion is liquid when warm and coagulates when cold. Some
say that the filling of the vesicles serves to excite sexual feeling,
but this is doul)tful in view of the fact that in some animals the
sexual desire exists before the vesicles are filled. Likewise, Stein-
ach found that rats, whose seminal vesicles had been removed,
still retained their desire for copulation. The function of the
secretion is to furnish much of the volume to the semen, and in
some way it has a distinct 1)earing on fei1:ility, inasmuch as ex-
tirpation of the organs in rats leads to lowered fertility. The
vesicles of the bull are in no sense a store-house for spermatozoa,
as is usually understood. Repeated examinations in a large num-
ber of bulls have led to the finding of spermatozoa there only in veiy
rare instances. That they serve as a resorption j^lace for sperms
that are not ejaculated is also very unlikely. Normally, one sees
on smear of the vesicles, occasional cells, leucocytes, lecithin gran-
ules, sago bodies, and rarely a few degenerated spermatozoa.

The CoLLicuLUS Se:mixalis is a rounded or cone-shaped emi-
nence in the posterior urethra, upon which the ducts of the sem-
inal vesicles and vasa deferentia open. The ducts open separately
at the bottom of two narrow slits, one on each side of the mound,
there being no distinct ejaculatory duct as in man. The function
of the eollicuius or veruinontanum is not definitely known. It
is generally believed that the structure is made up of blood spaces
which become engorged during erection, causing a l)lockage of
the posterior urethi'a. ^»•hich jirevents regurgitation of the semen.
Iivtina (27), howcNcr, demonstrated tliat the structure is not com-
posed of any unusual number of blood vessels or spaces, and that
removal of the organ Avas not followed by regurgitation of the
semen into the l)ladder during ejaculation. Tie ])elieves, and quite
logically, that its function is to afford a prominence upon which
the duets may empty. The mixture of the thick gelatinous semen
with the tliin ])rostatic secretion must occur at the moment of
ejaculation and must be jierfectly homogeneous, otherwise large
num])ers of the organisms remain in the thick gelatinous portion
of the fluid. The eminence serves this purpose in that the pros-
tatic ducts which converge toward it, may eject their secretion

15

toward the eminence, producing an admixture more evenly and
quickly.

Prostate: The hull possesses -what Ellenherger calls a diffuse
prostate. That is, there is no distinct glandular hody as in man.
It is composed principally of a glandular sheath around the
urethral wall. Just posterior to the neck of the bladder, and in
front of the urethral muscle, there is formed a slight dorsal trans-
verse elevation, extending downward on the sides. This is what
might be termed the body. The greater part of the gland is
"disseminate" in form, being a sheath of glandular tissue em-
bedded in the urethral wall. Dorsally it is about ten to twelve
millimeters thick, and ventrally about two millimeters. The gland
is a branched tul)ular structure, the interlobular tissue of which
contains much unstriped muscle. The lobules arc lined by a
columnar type of epithelium. The ducts, about thirty to forty
in numl)er, o])en into the urethra in two rows posterior to the
colliculus. The secretion is a thin, slightly turbid fluid, of a
faintly alkaline reaction. Its function is to dilute the semen,
stimulate the motility of the spermatozoa and nourish them.

Fish (28) believes that the activating property of the secretion
is due to enzymes, because boiling deprives the fluid of its power
to accelerate the motility of the spermatozoa. Serrlach and Pares,
quoted by ]\rarshall (29), working on dogs, have adduced evidence
indicating that the prostate is an internal secretory gland which
controls the testicular functions, and regulates the process of
ejaculation. It is stated that if the prostate is removed, sperma-
tozoa are no longer produced in the testis, and that the secretory
activity of the accessory genital glands ceases. These changes,
however, 'can be prevented by the administration of extracts of
the prostate. The fact that the prostate elaborates a secretion
having a definite relation to the testis, has been verified by other
authors.

Cowper's Geaxds (Bull)o-Urethral) : These glands are paired,
oval structures about one and one-half by two and one-half centi-
meters in size, situated on either side of the dorsal pelvic part of
the urethra close to the ischial arch. They are deeply embedded,
with the bull)us urethae, in the bulbo-cavernosus muscle, thereby
being inaccessil)le to palpation. Tn general, they are of a well
developed anastomosing tubular type. The connective tissue
.stroma is relatively thin, and thickens only in between the larger
lobules, where one finds the larger collecting ducts. Each gland
opens by a single duct. The epithelium is of the sim])le cuboidal
type, ijittle is known of the function of its .secretion, though
Kingsbury '':"50^ regards it in the light of a mating gland; that is.
it lubricates the genital passages dui-ing coitus, as does its
homologous structure in the female, Bartholin's gland.

Ellenberger describes the urethra, .slit ventrally, as presenting
the following picture: "The colliculus seminalis distinctly ap-

16

pears as a process or offshoot of the crista urethralis of the Tri-
gonium Vesicae. At the summit, and at the bottom of the two
slits, open laterally the ducts of the vesicles, and medially the
ductus deferens .... From the caudal slope of the colliculus
go two distinct mucous membrane folds which run through the
entire pelvic urethra, near together, somewhat diverging, and then
coming together, so that they form an elongated, narrow oval. At
their caudal union, the excretory ducts of the bulbo-urethral
glands open side by side. At the point of departure of the folds
from the colliculus, arises a niche-shaped opening, between both
folds, and likewise lateral to each fold. In these niches open the
ducts of the prostate. The openings of the disseminate prostate
lie in rows as in the horse, but form not less than six rows. There
is one row medial to each fold, and two lateral. Midler mentions
only the medial rows. These rows extend to the opening of the
ducts of Cowper's glands. The stratum glandulare (disseminate
prostate) is very easy to recognize with the naked eye."

Semen : The semen is the mixed product of the testicles, their
excretory passages, and the accessory sexual glands, a fact which
complicates its study considerably. The freshly ejaculated fluid is
cloudy, tenacious, more or less coagulable, and is rich in albumen.
It is weakly alkaline in reaction, and contains eighty to ninety
per cent of water. Of the solid constituents, there is forty per
cent of ash, of which three-fourths is calcium phosphate. Besides
the spermatozoa, the semen frequently contains epithelial cells,
leucocytes, concentric amyloid concretions, and lecithin bodies.
When' cold, the characteristic phosphoric acid salts are precipi-
tated. The fluid content is the product of the tubules of the
testicles, their excretory ducts, and the accessory sexual glands.
The characteristic odor "of the semen is supplied by a slimy nucleo-
albumen "spermin" which forms the spermatic crystals, and is
furnished by the prostatic secretion.

During ejaculation the spermatozoa and secretions added by the
testicle and epididymis are probably carried to the ampulla by
peristaltic muscular action, in the earlier stages of the orgasm.
At the height of the orgasm, the ampulla is emptied into the pos-
terior urethra in common with the secretion of the contracting
vesicles, here to be admixed with the thin prostatic secretion. The
entire mixture is then propelled, and ejaculation produced by
strong muscular contractions of the entire urethra. As was stated
before, the semen is the product of the testicles, the excretory
ducts, and the accessory sexual glands. The testes furnish the
essential germinal elements, the sperms, and some of the fluid con-
tent. Then is added the product of the epididymis, vas, and
ampulla, which stimulates the spermatozoa to active motility,
nourishes the organisms, and adds somewhat to the fluid bulk of
the secretion.

Stigler (25) states: "During its sojourn in the epididymis, the
properties of the spermatozoa are modified; the r.^vtility, ability

17

to resist heat, and other properties are augmented, at least in the
case of the guinea pig, rat, and mouse." To the secretion is then
added the product of the vesicles, Avhich contributes markedly to
its fluid content, nourishes the sperms, and supplies the ferment
which induces clotting of the semen when ejaculated. This is
very important because the clot formed in the vagina protects
the delicate spermatozoa from the hostile acid vaginal secretion.
The prostate likewise adds bulk and nourishing substances, besides
stimulating the spermatozoa to fuller and more lasting motility.
The addition of the spermin is perhaps unimportant. The func-
tion of the secretion of the Cowper's glands, which is added at this
time, is problematical. It does, however, have a diluting action on
the semen. Perhaps its secretion is poured out prior to ejacula-
tion so as to lubricate the canal and prepare the way for the semen.
Fish (28) has demonstrated by means of darkfield illumination,
the presence of numerous minute particles or ultraparticles in this
fluid. Their character and significance are matters of conjecture,
but it would seem as though they were not identical with the
"chylomicrons" or fat particles found in the blood by Gage. Per-
haps further researches wall reveal some intimate connection be-
tween the number present in a field, and the relative potency of
the animal.

Each portion of the tract furnishes some essential element to
the mixed product which is so remarkably adapted both as ^ a
vehicle for the ejaculation of the spermatozoa, and as a fluid in
which their motility is initiated and maintained. Any derange-
ment of one part is fraught with danger to the existence of viable
spermatozoa, and the continuation of full fertility on the part
of the animal. The physiology of each contributing gland must
be borne in mind at all "times. AValker (31) investigated the fer-
tility of the semen of the dog, taken from various parts of tho
tract. His results were: (1) semen from the testicle and head of
the epididymis showed no motility, (2) semen from the tail of
the epididymis showed some motility in the more fluid contents
of the preparations, (3) semen from the vas deferens appeared
about the same, (4) a mixture of epididymis semen and pros-
tatic secretion showed active motility, and (5) likewise in a mix-
ture of epididymis semen, though only in thase places where the
fluids had become well mixed. 'My observations, however, differ
in one respect with regard to the Inill, as I have found full motility
of the spermatozoa from the epididymis, but it is not so lasting as
when augmented by the addition of the prostatic fluid. Boettcher
(32) concludes: "that the secretion of the accessory male genital
organs possesses a protective colloid, which (1) hinders the sper-
matozoid action of the vaginal secretion, at least until the sperms
have time to reach the interior of the uterus which is an alkaline
reaction, (2) that it makes the ejaculate more voluminous, so that
by cohabitation, a very good part of the vagina becomes moistened,
and the spermatozoa become distributed over the greater part

2

IS

of the vaginal mucosa. This distri1)ution is rendered necessary
because some of tlie fluid flows from the vagina following coitus.
In this manner the opportuniiy is given for a part of the ejacu-
late containing the spermatozoa to be brought easier to the ex-
ternal OS, and (3) it happens that because of its content of sodium
chloride the life of the spermatozoa is stimulated and prolonged."
A fuller discussion of the essential physiology of the various parts
of the tract on the semen content, and fertility', Avill be taken up
later. The changes in biochemical content, reaction, and the re-
sult of the addition of bacterial products will also be more fully
discussed.

Spermatozoa : The history of the discovery of spermatozoa
is very interesting, and for that reason a brief outline will be
given. The "semen threads" were first observed in the year
1667, bv Ham, a student of Leeuwenhoek at Levden. The dis-
covery was announced, confirmed by findings in the dog and
rabbit, and discussed by the latter author under the title : Ob-
servationes Anthonii L. de natis e semine genitali animaculis
(Upon the formation of young from procreative material). The
sperms were taken to ])e animals on account of their motility, and
their significance remained questionable if not unknown. Spal-
lanzi, quoted by ^Marshall (29), was the first to show that the
filtered fluid was impotent, and that spermatozoa in the semen
were essential to fertilization. Kolliker, in 1841. discovered that
the sperms arise from the cells of the testis, and Barry in 1843.
observed the conjugation of sperm and ovum in the rabbit. This
led to a clear understanding of the function of these important
germinal elements.

The spermatozoa are the male procreative cells, and are cliar-
acterizecl by the possession of a head containing the chromo.somes
necessary for fertilization, and a tail capable of propelling the
organism on its way to meet the ovum. The length of the entire
sperm, including the head, is seventy-five to eighty microns. The
head is nine and five-tenths microns long, and five and five-tenths
microns wide. It may be divided into two principal parts, the
head and tail. The head, for the larger part, is made up of the
nucleus, and may be differentiated by staining reactions into a
darker staining posterior part, an anterior lighter part, and often
a still lighter area between the two. On the anterior part is a
sharpened edge, the acrosome, which serves to perforate the ovum.
The Avhole is surrounded by a very definite limiting membrane
which often becomes obscured under abnormal conditions. The
tail may be divided into three parts: connecting piece, principal
part, and terminal filament. The connecting piece, the essential
motile apparatus, is the thickest and strongest part, and joins the
tail p-roper to the head. It consists of the central axial filament, a
spiral filament around this, and an outer mitochondrial coverinsr.
Anteriorly it is limited by the anterior portion of the posterior
chromosome, and posteriorly liy the annular chromosome. The

19

anterior chromosome is directly connected with the head, there
appearing a light break here at the neck where separation fre-
quently occui-s. This neck ser^-es as a joint for the motion. The
axial filament, therefore, does not reach the head, but extends
back from the anterior part of the posterior chromosome. The
principal part consists merely of the axial filament, and a thin
outer covering, while the end piece is quite thin and is made up
solely ol the uncovered axial filament. The finer structures are
seen only when special staining reactions are used, and then only
when the sperms are obtained directly from the testicle. The
function of the sperms is of course primarily that of fertilization.
Numerous observers have, however, thought that they might have
some other definite, thougli unknown, use.

An editorial in the Journal of the American Medical Associa-
tion (33) raises several important questions regarding this ob-
scure phenomenon. The fact that an enormous number of spenna-
tozoa are produced, and only one, or at mo.st, a few perform the
function of fertilization, raises the question as to what becomes
of the remainder. It is stated: "Zoologists have found that in
some of the invertebrates the spermatozoa invade the entire body
of the female, and in some species they reach the ovum by pene-
trating the cuticle from outside and migrating to their goal.
Studies on rodents show that the sperms invade the epithelium of
the generative mucosa and underlying connective tissue. These
tissues seemed to be stimulated to growth, suggesting that this
may influence the uterine mucosa in its preparation for receiving
and embedding the egg, and in forming the decidua." It has
been shown that the sperms contain a specific protein capable oC
producing antibodies in the blood plasma, by citing the fact that
rabbits develop a distinct Aberhalden reaction for testicular pro-
teins shortly after cohabitation. One very important observation
showed tliat by immunizing female rabbits with sperms they were
rendered sterile for some time, although after a few months they
again became capable of impregnation. The question raised is:
**.... if the spermatozoa invade the female tissues and cause
the formation of specific antibodies Avhich are capable of prevent-
ing fertilization, may not such a process participate in the prob-
lem of sterility?" This very problem seems to be a factor in ex-
plaining why some couples who are not fertile to each other
subsequently are both fertile when they cohabit with other indi-
viduals.

Motility: After clinical observation of the motility of the sper-
matozoa of the bull, I find that it differs little or none from the
types as observed by Eeynolds (34) in his work on human sperma-
tozoa. His ol)servations are so accurate and well described that
they will be given in his words. "All normal motions appear to
be consecutive phases. Initial motion, i. e., motion as seen in
fresh semen under favorable conditions, consists of a lashing
of the after part of the tail from ^ido to side which is so lapid

20

as to constitute vibration. It produces rapid forward motion in a
practically straight line, the head, middle piece, and forward por-
tion of the tail maintaining their position in the line of motion
with practically no swaying from side to side. The action of the
flagellum is so rapid that it is quite impossible to follow its indi-
vidual movements. Spermatozoa swimming in this manner head
against the current and usually cross the field of observation in
about five seconds in the absence of currents or obstacles. This
type of motion will be described throughout the paper as 'pro-
gi'essive vibratile' motion. Progressive vibratile motion is nor-
mally succeeded after a variable length of time by what I regard,
as the second phase of normal motion.

"The second normal motion differs from the first not only in
its character but in markedly reduced speed. The tail movement
alters to a long stroke from side to side and almost the whole
length of the tail partakes in the stroke. This is, moreover, accom-
panied by swaying of the head and middle piece through an arc
which is always considerable and may even equal ninety degrees.
The general outline of the spermatozoa, from being practically
straight with almost non-detectable sharp, quick, small arc vibra-
tion of the aftertail, has become an S in outline, with large, slow,
plainly perceptible undulations traveling gradually backward
throughout the length of the spermatozoon. Speed has been lost
and direction seems to be more specifically determined by the
surroundings. Individuals at this stage show a pronounced choice
of direction and go up to objects in the medium, from which they
later make off as though the movement were determined by tactile
reaction to some extent. This type of motion has, therefore, been
named 'undulatory tactile' in contradistinction to 'progressive
inbratile. '

'The third type of normal motion succeeds the second and con-
sist.s in a tendency on the part of the spermatozoon to push itself
against or into any small masses of cells, or sometimes other
materials, which it may find in the neighl^orhood, bunting itself
into any small cove that can be found, and maintaining a slight
burrowing motion by a lashing tail movement of the vibratile type
not unlike the movements of the caudal fin of a fish. The move-
ment of the flagellum in this third type is unlike the second type
in that it is wlbratile rather than lashing, but is slower than the
\nbratile motion of the first type and less limited to the after-
part of the tail. The?e spermatozoa are apparently not caught
in the debris or unable to move off. From time to time, they
back out of such a cove and seek another mooring place.

"This 'stationary bunting' motion is less universal than the
other two. Many individual spermatozoa fail to attain it. It
seems probable that only the most vigorous individuals ever reach
this stage. It has not been encountered in unmixed semen or in
any artificial mediums. It has been observed only when the sper-
matozoon is in the secretions of the female genital tract. It is

21

most frequent Avhen the spermatozoon is in contact with a nest of
epithelial cells ....

"The three types of normal motion are not only distinctive but
are always consecutive, i. e., the second follows the first after
a period which is apparently determined both by initial vitality
and by the favorable or unfavorable character of the medium,
while the third has been observed to occur only in individuals which
have already developed the second. They apparently constitute a
normal cycle.

"This cycle is open to the theoretical explanation that these
types of motion are directly- adapted to the function of the sper-
matozoon ; thus, the progressive vibratile motion which is char-
acteristic of the earliest period of its existence appears especially
suitable for its prolonged journey through the cervix and uterus
to the fundus and tube. This is supported by the fact that during
this motion it always heads directly against any existing current,
and that during this stage of its journey it must under natural
conditions be continuously exposed to the faint outward ciliary
currents of the mucous membrane of these passages.

"The undulatory tactile motion which succeeds the progressive
vibratile would then be well adapted to the later stage in which
the spermatozoon has reached the tube, where its success in conju-
gation is dependent on its finding the ovum rather than on further
progress.

"The stationary- bunting type of motion is that which would be
demanded by the passage of the spermat07X)on through the egg
membrane which has been so often observed in the lower animals.
This very plausible explanation is, however, necessarily theoretical
and must always remain so, as the conditions which surround the
specimen on the field of the microscope vary in so many respects
from those in which it accomplishes conjugation in the course of
nature; but the practical importance of the study of tj^pes of
motion is not affected by their explanation."

The duration of motility is a variable factor, dependent entirely
upon the environment in which the spermatozoa are placed.
Within the body they usually sur\'ive at least a week. One author
describes a case in which he found living sperms in a Avoman who
stated that coition had not been experienced for three and one-
half weeks. It has been stated with regard to human sperms, that
their motion should continue or be cai)able of being re-established
for twelve hours. Gary (35) states, "first, that in their proper
medium and at the body temperature the viability of the sperm
cells may extend over a period of a few days ; second, that their
prolonged vitality is probably dependent upon the normal lime
salts of the prostatic fluid ; third, that the sustaining power of
the seminal fluid is increased by its undon with the normal secre-
tion of the female genital tract." After death of the male animal
they retain their motilitj' in the genital tract for twenty-four to
forty-eight hours.

22

Wolf (36) worked on this problem in rabbits, and summarizes
as follows: "The motility of rabbit spermatozoa can be preserved
for at least nine days by placing the juice of the epididymis in
the Tyrodc solution which had been buffered and to which glucose
had been added. The solution is adjusted to a pH of about 7.4.
Oxj'gen is passed in and a suitable amount of sodium bicarbonate
added. The preparation must be kept at a temperature near the
freezing i^oint of water." Under ordinary conditions motility
persists outside the body only a few hours after ejaculation, but
if the semen is kept quite cool till the time of examination on a
warm stage, motility should be capable of being restored in at least
a fair percentage of the sperms for twenty-four to forty-eight
hours. The cells are more sensitive to heat than to cold, and even
to dilute acids more than to alkalies.

Henle (quoted by Ellenberger) states that a spermatozoon under
favorable conditions travels at the rate of twenty-seven millimeters
in seven and one-half minutes, which makes three and five-tenths
millimeters per minute. This is about sixty times the entire
length of the spermatozoon, and twenty-one centimeters in an
hour. Forward motion is also more pronounced when the swim is
against the current, such as is produced by the cilia of the oviduct.
It has been demonstrated that feebly motile sperms become very
actively motile when placed on the mucosa of a fresh Fallopian
tube.

Technique

The material used in the work came from abattoir animals, bull
calves and adult bulls raised in the department herd, and sires
upon which clinical observations had been made by various veteri-
narians in the field. Senlen samples, many of which were sent in,
were collected as often as possible after the method described by
AVilliams (16). The genital organs were removed with as little
chance of contamination as possible, and taken or sent to the
la1)oratory where the examinations were made soon after arrival.

All cultures were made by searing the surface carefully, tearing
out a small portion of the tissue with sterile forceps, and placing
it upon the media. In most cases, however, where fluids were
])resent, tubes were inoculated with the material which had been
drawn off with a sterile pipette. As stated by Carpenter (9), in
his work on the female genital tract, the organisms usually live
in the depths of the tissue. The media used principally were glu-
cose glycerin agar (glucose 1 per cent, glycerin 3 per cent) ; plain
agar, both with a pH value of 7.4, and Loeffler's blood serum.
Small amounts of sterile blood serum or defibrinated blood were
added to most of the agar slants to insure better growths of
streptococci when present. All tubes, to which the serum had
been added, were incubated for forty-eight hours before inocula-
tion to insure absolute sterility.

After inoculation, the agar tubes were sealed with sealing wax
to give a partial oxygen tension which was quite necessary in iso-

23

lating the streptococci. The growth of other organisms was by
no means hindered by the procedure, for one tube from eacii
organ was often left unseaKxl. Incubation was at ol'^ C, and the
routine method of examining the tubes was identical with the
method of Carpenter (9).

Whenever possible, a sample of blood was obtained from the
animal either before, or at the time of slaughter, for agglutination
wuh Bact. aburtum antigen. Extracts from the semnial vesicles,
testes, and epididymes were injected into the guinea pigs and
examined at the end of four to six weeks for the presence of
Bad. ahortum.

Sections of all organs were fixed as soon as possible in either
Zenker's or Ilelly's fluid. Hematoxylin and eosin were used as
routine tissue stains. Eosin and methylene blue, and Mallory's
connective tissue stain Avere, however, frequenth' utilized for
special staining reactions.

The motility of the spermatozoa is best observed about half an
hour after ejaculation, when the thick tenacious clot has started
to liquefy. A drop of the fluid is placed upon a warmed slide,
preferably one with a slight depression in it. and observation
made with high or low powered objectives. The semen may be
examined whole, or diluted with physiological saline .solution. In
the latter ease, the sperms have a greater opportunity for freedom
of motion in the absence of the thick viscid coagulate. A small
vial erf saline solution may be carried in one's pocket where it
will be kept warm, and a drop of this placed upon the glass slide.
If the clot of semen is merely touched to this drop on the slide,
plenty of .spermatozoa will be deposited for an examination. This
method is very satisfactory for the observation of motility, but
needless to say, the undiluted semen must be used for the deter-
mination of the number of sperms present. If necessary, the
specimen may be covered with a cover glass and the oil immersion
objective used. While a warmed slide is quite sufficient to enable
one to detect the i^resenee of motility, the field soon cools and the
sperms gradually become less motile. If possi])le. it is best to
use a small electrically heated stage warmer, which keeps the
field at a constant ])ody temperature, so that the duration of the
motion may be observed for hours if warmed jihysiological saline
solution is added as the fluid evaporates.

Stained preparations are best made with fhin smears on the
glass slides. This is conveniently done by placing a drop of the
semen on a slide and smearing it over the surface with the edge
of another slide. A fairly thin and even field is thus obtained. A
.still better method is to first dilute the semen with physiological
saline solution, so as to ol)tain fewer sperms in the field. After
drying the pre]>aration in air. fixation may l)e produced by draw-
ing the slide through a gas flame several times, by immersion in
equal parts of alcohol and ether, or even 1)y the use of tissue fixers
such as Ilellv's or Zenker's .fluids. For ordinary .staining, heat

24

fixation is the quickest, and at the same time is quite satisfactory.
After the slide is cooled, or washed, to remove the fixing solutions,
it should be placed for a few minutes in a freshly prepared solu-
tion (1 per cent) of chlora/^ene, as recommended by Williams, to
remove the mucus and proteid material which otherwise blur the
field. Other authors (38) have recommended diluting the semen
with about twenty volumes of a 0.12 per cent sodium carbonate
solution in 0.8 per cent sodium chloride. From this liquid the
cells should be centrifuged for several minutes, removed with a
pipette, and smeared on the slide. Following this, the slide should
be thoroughly washed, preferably for ten minutes in running
water, after which it is ready for staining. Numerous methods
have been used for this procedure, but the sperms are more or
less erratic in their reactions to the dyes, and one must be very
careful to use the same method in all samples, in order to obtain
uniform results. For quick staining, to bring out gross abnormal-
ities of structure, and number of sperms present, one may use
Gram's stain, or a light stain with any of the aniline dyes, such
a.s fuchsin. To bring out the finer structure, particularly of the
head, more careful technic must be employed.

Carnett and others (38) recommend the following: " The
method of staining by iron-hematoxylin, particularly when supple-
mented by a cytoplasm stain, has proved, on the whole, the most
satisfactory, and possesses the additional advantage of being abso-
lutely permanent, a quality that few anilines can boast. The
method consisted of treating the fixed object — and here the fixing
agent was heat — with a two per cent solution of iron-alum for
from two to four hours. The excess of iron-alum was then com-
pletely removed 1)y pure water, and the object treated with a solu-
tion of hematoxylin (one per cent aqueous) for twelve hours or
longer. The cells by this time were perfectly black. However, a
1 per cent solution of iron-alum removed the stain from the cyto-
pla.sm. leaving the chromatin of the head, the centrosome, and the
axial filament a brilliant blue-black. Care must be taken that the
preparation is not over-decolorized. After deeolorization a satu-
rated aqueous solution of eosin was added for from one to three
minutes. This stained the protoplasmic envelope pink, and, unless
the envelope is overstained, the view of the inner structures is not
impaired in the least."

AVilliams (17) recommends using two staining solutions, one of
alcoholic eosin and fuchsin, the other a diluted methylene blue.
The results obtained are, however, more or less erratic, due to
the unstable character of the former stain, and the ease with
which one may over or under stain. Many beautiful specimens
may, nevertheless, be obtained by this method. I have frequently
used a fairly quick method, though one not satisfactory in ail
cases, which consists of staining for five or six minutes in a satu-
rated aqueous solution of fuchsin, washing in water, and counter-
staining for a few seconds in a strong solution of methylene blue.

25

A quite satisfactory method is to stain from two to five minutes in
a saturated aqueous solution of methyl green, with the application
of gentle heat. The heat may be applied by warming the slide over
a gas tiame as it steams, or by placing the jar containing the stain
in a hot water bath. The slide is then washed thoroughly and
eounterstained for five minutes in a strong aqueous solution of
eosin. This is a fairly reliable method, and many excellent pre-
parations may be obtained by its use. The nucleus is stained
green, the anterior part of the head and all of the tail pink. So
far, I have found this a very reliable stain for routine work.

Pathology

In the genital tracts that I have studied, a complete pathological
and bacteriological examination was made Avherever possible, but
in man}' of the abattoir animals, and certain others, gross and
microscopical examinations only could be made. The genital
organs of one hundred and ninety-six males have been examined,
and the gross or miscroseopical changes, or both, determined. The
abattoir animals Avere from a large slaughter house, and a small
local plant.

Of the tracts, the pathology of which was studied, two were
from aborted fetuses ; seven from apparently normal young calves ;
four from mature fertile bulls ; and sixteen from mature infertile
or sterile sires. The remainder (167) of those examined were
from abattoir animals. Besides these, three specimens of seminal
vesicles, and seven of testes were studied.

The tracts of the aborted fetuses and veal calves were apparently
normal, both on gross and microscopic examination of the vesicles
and testes. On gross examination, the tracts of the mature fertile
bulls were normal, except for the presence of many fine connective
tissue tufts and strands upon the serous covering of the tails of
the epidid>Tnes, and adjacent portion of the parietal layer of the
tunica vaginalis. jMicroscopic sections of all parts were apparently
normal. The more important pathological changes in the tracts of
the sixteen sterile or infertile bulls are given in the appended
chart. The tracts are numbered the same as in Group VI of the
report of the bacteriological findings ; that is, any particular num-
ber in either tal)le refei's to the same animal. Eeferences are made
throughout the text to some cases which appear in this group of
animals. Prostate and Cowper's glands are not included in the
chart as they were not examined in some, and were negative in the
others. Fibrous tufts and strands were present on the covering
of the epididymes in each animal.

The study of sections from the abattoir animals, as well as those
from the sterile or infertile bulls, forms the basis for the follow-
ing observations upon the pathologA' of the male genital tract.
The tracts secured from the abattoir were studied for the most
part on the ba.sis of the organ rather than on that of the animal.
For example, all sections of testes were placed in the same bottle
of fixer, and the same plan followed for the other organs.

26

Testes : The testes seldom presented gross alterations of struc-
ture except for abscess formation, which, according to Williams,
occurs more frequently in the bull than in any of the other
domesticated animals. He also states that arrest in development by
which the organs remain soft, flaccid, and somewhat smaller than
normal is not uncommon. One very interesting specimen, which
typifies abscess formation, came from a bull with a history reveal-
ing that one testis had become much enlarged, hot, and painful.
These sj^mptoms developed very rapidly. Anorexia was well
marked. Local applications were used for several weeks, but
at the end of two months the condition was so little improved that
unilateral castration Avas performed. The general condition of the
animal soon improved, but after a year of service he was so uncer-
tain compared to what he had been, that he was sent to the
butcher. It was impossible to obtain the other testicle for study,
though it undoubtedly was abnormal. The testicle removed was
considerably enlarged, measuring twenty by ten and one-half centi-
meters. The tunica albuginea presented a thickness of six milli-
meters, and was made up of firm sclerotic tissue. The epididy-
mis was not recognizable in the mass. Testicular tissue was almost
entirely gone. The only remains, of what appeared to have been
I)arenchyma, was an elongated irregular area at one side of the
organ. This tissue consisted of a whitish opalescent material,
speckled with varying sized abscesses. This organ is pictured in
Fig. 3. The remainder of the organ consisted of a thick yellowish
caseous mass. Strepiococcns viridans was recovered from the
outer portion of the organ, and guinea pig inoculations failed to
demonstrate Bact. ahortum.

^Microscopically, changes are quite common and varied in char-
acter. In the seminiferous tubules, the changes range from a
slight desquamation of the germinal epithelium to atrophy and
complete degeneration of the entire tubule, as was the case in the
left testis of Bull 1. In the mild cases, spermatogenesis occurs
apparently in a normal manner up to the spermatid stage, at
wliich point many of the cells degenerate and slough off. These
appear in the seminal fluid, associated with the few sperms that
reach maturity. This sloughing and degeneration may be localized
in a few of the tubules, or it may be widespread over the entire
organ. Likewise, the changes may involve not only the more
mature cells, but they may be so .severe as to cause almost total
degeneration and desquamation of the seminal epithelium, as in
Fig. 15. These defects in spermatogenesis are of course evidenced
in the semen l)y the presence of immature, or abnormal types of
sperms. With cessation of spermatogenesis or degeneration of the
epithelium of the entire gland, no sperms are formed. Not infre-
(inently one finds numerous tubules, or even the entire testis in
which the germinal epithelium is intact, ])ut there is little or no
evidence of mitosis, as in some tubules of Bull 6. The cells are
several layers deep as in the normal condition, but they are not
dividing. This condition is shown in Fig. 13.

27

In the more chronic forms, the tubules become atrophied, and
frequently disappear entirely. The membrana propria may be-
come thickened, due to excessive connective tissue formation, or
infiltration with serum or exudate. On the other hand, a dis-
tinct atrophy may occur. The sti'oma of the organ not i)ifre-
([uently is thickened by inflammatory exudates, or by a noticeable
increase in the connective tissue. In some testes, the connective
tissue is so much increased that the tubules rapidly become
atrophied, and disappear. In abscess formation, due to acute in-
flammations, the entire organ Ijecomes enlarged, markedly hyper-
aemic, and infiltrated with leucocytes. Necrotic areas appear here
and there in the parenchyma. The rete often shows a marked de-
generation of the lining epithelium, and atrophy caused by in-
crease of the interstitial connective tissue.

Epididymis: This organ not infrequently presents gross ab-
normalities, and very often is pathological on microscopic examina-
tion. Acute inflammation, with induration or abscess formation, is
very common in the tail, but less so in the head and body.
Possibly this is caused by the fact that the tail is the most pendant
portion of the organ. In these cases, the tail is enlarged, soft, and
quite hot and painful on physical examination. Enlargement, due
to a connective tissue induration, occurs occasionally in all three
parts, and the inflammation may produce adhesions to the adjacent
serous membranes. Inflammation of both the parietal and visceral
layers of the tunica vaginalis is very common. In those cases,
the membrane usually is quite hyperaemic. and on the surface it
presents many small reddened tufts of newly-formed connective tis-
sue. In adult bulls it is exceptional not to find at least slight evidence
of some previous inflammation. In all of the numerous bulls ex-
amined, both apparently normal and sterile, I have found but one
in which some evidence of inflammation (either present or past)
could not be found. Along with the fibrous tufts, are numerous
fine strands of connective tissues passing from the covering of the
tail of the epididymis to the adjacent portion of the parietal layer
of the tunica. The strands often extend even to the upper part of
the head.

INIicroscopically, inflammation of the part is shown by hyper-
aemia, loss of cilia of the lining cells, and exudation. In the more
.severe forms, the lining cells which furnish considerable secretion
for the nourishment and stimulatioji of the sperms, l)ecome de-
generated, and are exfoliated into the lumen, as in Fig. 21. This
condition is very common in sterile bulls, and those of lowered
fertility. In the chronic types, the interstitial connective tissue is
increased in amount, leading to degeneration and atrophy of part
or all. of the tubules, as in the case of Bull 2. Infiltration with
leucocytes, and necrosis, are the predominating lesions in the p.vo-
genic types of inflammation.

28

Ductus Deferens : This tube seems to be peculiarly free from
severe inflammatory processes, and when these appear they are
limited to the mucosa. The cells of the lining membrane not in-
frequently show a mild type of degeneration and exfoliation, or in
the more chronic forms, the entire membrane degenerates and disap-
pears. In man, the duct occasionally becomes occluded, but so far
I have failed to find this condition in the bull. Undoubtedly, when
the occlusion does occur, it is near the origin of the duct at the
tail of the epididymis.

Seminal Vesicles: The seminal vesicles and epididymis,
especially the tail, seem to be the parts most subject to extensive
pathological changes, and bacterial invasion. In most instances,
diseased vesicles present gross manifestations recognizable on
clinical examination, while on the other hand microscopic changes
may be present in the absence of gross lesions. As diagnosed on
physical examination, or even on post mortem examination of the
tract, the various forms may be classified into :

1. Acute Catarrlial Type: In this form, the vesicles are usually
enlarged, soft, and more or less reddened by hyperaemia. On
physical examination of the animal, distinct flinching is produced
when pressure is applied to the organ. Enlargement may even
be absent in the early stages, and the diagnosis may be made from
the extrenie sensitiveness alone.

2. Suppurative or Cystic Types: In both of these types, the
vesicles are usually enlarged, either uniformly, or, as is usually the
case, in localized areas. The suppurative form may extend over
the entire gland, forming one large encapsulated abscess, or on the
other hand, it may take the form of variable sized abscesses with
thick sclerotic, or thin fluctuating walls. Occasionally the abscesses
rupture and discharge their contents into the rectum. Dr. Wil-
liams presented one case of this type. One vesicle was apparently
normal, whereas the other was about five times larger than normal,
and consisted of a dense outer capsule which was adherent to all
surrounding parts. On dissection, it was found that the organ
consisted of abscesses of various sizes, the larger one of which had
ruptured some time previously into the rectum, leaving the dis-
tinct remains of an opening into that part. The cystic form may
occur either with or without suppuration. One case came to my
attention in which both vesicles were made up of abscesses of
varying sizes as well as of a smaller number of cysts. Evidently
the cysts were of the retention type, and were secondary to the
pyogenic infection,

3. Chronic or Sclerotic Type: This form is characterized by a
distinct firmness with or without marked enlargement. The con-
dition may be accompanied by disease of the parenchymatous tissue
or it may take the form of a chronic productive inflammation of
the interstitial tissue. This inflammation may be simply a super-
ficial thickening, or it may extend in between the lobules.

29

4. The Peri-vesicular or ^'pan-inflammatory" Type usually is
the result of severe inflammation of the vesicles, with probable rup-
ture of some of the smaller cysts or abscesses upon the surface.
The vesicles are, as a rule, considerably enlarged and buried in a
dense mass of adhesions which involve neighboring structures.
The vesicles cannot be palpated on i)hysical examination, and it is
only on careful post mortem dissection that they may be studied.
This type, however, is quite rare, — two cases only having come to
my attention. In both, the vesicles themselves were markedly
affected.

Microscopically, changes in the vesicles are quite frequently en-
countered, even in the absence of gross manifestations. In the
•acute catarrhal forms, the mucosa and submucosa are hyperaemie.
The lining cells show various forms of degeneration, and there
are, as a rule, inflammatory exudates in the lumen. As the in-
flammation progresses, the lining cells degenerate further, and
become cast off into the lumen of the glandular cavities, as in
Plate VI. The normal clear mucous secretion becomes mixed with
fibrin, leucocytes, and cellular debris. These changes may involve
merely parts of the organ, or they may be quite extensive. With
large sections, one may find the inflammation in all stages, from the
mildest catarrhal type, to complete degeneration and exfoliation
of the secretion-forming mucosal cells, and filling of the cavities
with degenerated cells, leucocytes, and debris. Frequently the inter-
stitial tissue is in no way affected, but at times it is thickened by
©edematous exudates, leucocjiies, and fibrin. The chronic inter-
stitial form is characterized by a considerable increase of con-
nective tissue, — producing marked atrophj', or even complete
obliteration of the glandular cavities. Microscopically the sup-
purative form may be diffuse over the entire gland, or as stated
previously, may be in the form of localized abscesses, with or with-
out a thick connective tissue Avail. The parenchyma in these cases
is usually extensively degenerated and atrophied in those parts
that have not undergone suppuration and necrosis. The cysts
appear to be of the ordinary retention type, and may or may not
be accompanied by extensive changes in the lining epithelium.

Both the abscess formation and cystic conditions are undoubt-
edly initiated by an obstructive inflammation of all or part of the
excretory duct. This is, however, a protective mechanism, for
Avhere the duct is closed the bacteria and exudates are unable to
reach the urethra and contaminate the semen.

Prostate and Cowper's Glands: These glands were more or
less neglected in the early part of the work, but later were sub-
jected to the same examination as other parts. Of the thirty-six
of each type of gland examined, I failed to find one with any gross
changes, l)ut two prostates were found that presented a mild
catarrhal inflammation of the mucosa. It is probable that Cow-
per's glands, as well, occasionally undergo inflammatory changes.

30

Semen : The semen, made up as it is of mixed products of
the testes and accessory sexual glands, is very often abnormal, as
would be expected in view of the frequency with which changes
occur in the glands contributing to its formation. The normal
semen is remarkably adapted to its function of nourishment and
stimulation of the sjiermatozoa, and their conveyance to the in-
ternal female genital organs. The spermatozoa are extremely
sensitive to changes in their environment, with the result that any
alteration of the physical or biochemical content of the seminal
fluid may cause death of the sperms. With this in view, we must
remember that disease of any of the contributing organs is a poten-
tial danger, and threatens the potency of the animal. Each or all
of the glands may add bacteria, acid secretions, or inflammatory
exudates. On the other hand, they may not function at all. In
each case, however, the semen is altered.

Unfortunately it is impossible with present methods to obtain
the fluid absolutely free from vaginal mucus, but with care it may
be secured reasonably free from contamination by douching the
prepuce of the bull and vagina of the cow before service. This
method was used as often as possible in collecting the samples.
The usual amount of semen obtained was from six to ten cubic
centimeters.

With a hj'persecretion of one or all of the glands, the semen
becomes quite thin and watery, with a deficiency of solid matter,
together wdth changes in reaction. On the other hand, hypofunc-
tion results in a secretion too viscid, which is equally unsuited to
the requirements of the spermatozoa. The thin watery semen
clots imperfectly or not at all, and clotting is essential in protect-
ing the spermatozoa from the acid secretions of the vagina.
Likewise, a medium too viscid is a distinct hindrance to motility.
Changes in reaction are very frequently encountered. The sperms
are very sensitive to dilute acids, so that with even a slight acidity
motility may diminish or entirely cease. Purulent inflammatory
exudates are occa.sionally mixed with the semen, and although the
pus cells themselves have not been found to be destructive to the
sperms, certain degeneration products in the exudate are very
toxic, and inhibit or destroy the motility. So far, I have failed
to find red corpuscles present. One very interesting sample of
semen was quite thick, of a yellowish green color, and of a dis-
tinctly acid reaction. The secretion from the vesicles was later
found to be of this same character, and was due to a Ps. pxjocija-
neus infection. The vesicles were highly inflamed and degenerated.
The spermatozoa were in this case markedly decreased in number,
and devoid of motility.

The early precipitation of the "Boettcherchen." crystals S€ems
to be intimately connected with sterile semen, or spermatozoa of
lowered vitality. Likewise, a decrease in solid matter is often seen
in a deficient secretion. In normal semen, the clot disappears after
standing a time, and a thick .sediment settles out. This sediment is
decreased in amount as a rule in abnormal semen.

31

Spermatozoa : Spermatozoa, the essential germinal elements,
are very frequently abnormal, changes in which may be manifested
in many ways. We may divide the deviations into changes in
structure, and changes in the motility which is so indicative
of the intrinsic vitality of the sperm. Reynolds (34) describes
two forms of abnormal motion. The first is " rotary swim-
ming," in which the sperms move forward progressively, and
sometimes with fair rapidity, but in a spiral screwlike manner, lie
states that this type of swimming is very awkward, easy to recog-
nize, and is usually of quite long duration. The other form termed
"pendulum swimming," he slates, is less common than the rotary
swimming and is usually confined to relatively fewer sperms in a
given field. "In this the middle piece and upper tail seem to lose
their flexibility and balance to a considerable degree, and the
lower tail motion swings the forward part of the spermatozoon to
and fro with a pendulum movement. This type oi: swimming yields
very poor progress."

One factor we must bear in mind in the study of the semen ob-
tained from the vagina, is that the spermatozoa may be highly
motile before ejaculation, but the admixture of hostile vaginal
mucus may inhibit or destroy the motility. On the other hand,
the conditions may be reversed. Gary (35), in one instance,
found that the spermatozoa in a sample of semen collected from a
condom, appeared to be of very low vitality, Avhile when thej^ were
mixed with the vaginal secretions, an (exaggerated activity was
manifested. INIay we not have to contend with this factor in some
herds in which there is a very distinct 'acid and toxic vaginal
secretion from the products of cervicitis and vaginitis'?

In a study of motility, we must consider not only the abnormal
types which may be encountered but the percentage of motile cells,
and the duration of the movement. In nccrospcrmia all the
ejaculated cells are motionless or dead. In other specimens, vary-
ing percentages of the cells are without motion, and the others may
be possessed of full and lasting motility. On the other hand, the
motility in some eases is veiy active at first, Init quickly subsides
even undin* the best of conditions. The appearance in freshly
ejaculated semen of numerous sperms that have a tendency to take
on the "undulatory tactile" type of motion when they should be in
a hitxhly active state, is very indicative of lowered vitality. Many
specimens present this very picture, whereas the very active pro-
gressive movement should, under proper conditions, survive for a
considerable time before it gives way to the second, and slower
type. The cells frequently enrly bunt into epithelial cells or cUimps
of immotile sperms, then back out and move around sluga'ishly,
only to repeat the same performance till they stop moving en-
tirely. I have seen one specimen in which the sperms all tended
to clump. Whether this was the result of some agglutinative sub-
stance in the vaginal secretion is problematical. I have seen several
specimens of semen in which practically all the sperms were motile

32

when first examined, but the motion did not sursdve for any great
length of time. Even a small percentage of motionless sperms or
of those sho^\^ng lowered vitality is a considerable factor in
potency. xVlthough millions of the germinal elements are ejacu-
lated into the vagina, large numbers of them are destroyed or be-
come motionless there, and a small number is left behind in the
cervix and uterus ; so that even though but a single sperm is
required for fertilization, the chances of impregnation are dimin-
ished in proportion to the number of dead or defective sperms.

Aspermia: Absence of spermatozoa in the semen is rarely en-
countered, and is probably due either to total cessation of sperma-
togenesis, or to an obstruction at some point in the system of
excretory ducts. I have seen but one case of this character. The
semen of this bull was greatly increased in amount, and of a thin
watery consistency. Due to lack of cooperation on the part of the
owners, the tract could not be obtained for study. Oligospermia,
or a diminution of the number of spermatozoa, is quite common,
and is undoubtedly associated with defective spermatogenesis,
either as a result of poor mitosis of the seminal epithelium, or
degeneration of the elements before maturity. This condition may
vary from the finding of only occasional dead sperms in the field, to
but a slight decrease in the usual number of normal sperms
observed.

Abnormalities in morphology may be classified into immature
types, and deformities or imperfect development of the head and
tail. Defective spermatogenesis occurs so frequently that it is
not surprising to find spermatozoa in various stages of develop-
ment cast into the excretory ducts. The various stages passed
through in the development, from spermatogonia to adult sperm, are
numerous, and it therefore is to be expected that we should see in
abnormal semen many different immature forms. No classification
of the various types can be made, but a clearer understanding of
them can best be obtained by a review of the process of spermato-
genesis.

Spermatocytes and spermatids are seen in the more severe types
of defective spermatogenesis, and are relatively uncommon, while
the more mature forms that result from the transformations of
spermatid to adult cell are very often seen. Some of these inter-
mediate types are large oval cells without distinct nuclei and as a
rule with poorly developed tails. Cells with no tails or distinct
nuclei, those with protoplasmic appendages to the head or tail, and
various other types, are occasionally encountered. IMost of these
are motionless and incapable of producing impregnation. Others
are active, but survive a comparatively short time. According to
Gary, the production of the immature cells is an effort on the part
of the testes to sunplv an abnormal demand, and their presence
indicates that the fertility of the semen is impaired.

The deformities, which may be divided into cephalic and caudal
groups, are also the product of defective spermatogenesis, or they

33

represent a degenerative process induced possibly by abnormali-
ties of the fluid environment. It is rather difficult, however, to
distinguish between deformities and immature types. The two
most common cephalic deformities are Avhat might be called macro
and microcephalic forms. In the former, the head is enlarged to a
greater or less extent, it is usually defective in staining qualities,
and its outline is indistinct, due to degeneration of the covering
membrane. This type is seen in Fig. 32. Also the shape of the
head is usually abnormal, being either quite rounded, loifg and
narrow, or short and very broad. Cells with protoplasmic ap-
pendages, though they are more properly an immature type,
occasionally give the head a greater volume. Microcephalic sperms
vary from those slightly smaller than normal to those in which the
head is represented by a slight knob. In some cells, the head is
small and round, in others, short and stubby, while another type is
normal in outline but diminutive in size. These forms likewise
are, as a rule, deficient in staining qualities, and are undoubtedly
degeneration forms, occurring either as the result of faulty de-
velopment, or degeneration subsequent to their formation. Gary
believes they are degeneration types because in the majority of cells
the tail is apparently fully formed, and in the normal process of
evolution the tail is the last part of the cell to be exhibited.
Double headed forms are quite rare, but they nevertheless appear
at times. Their significance is difficult to explain. Another very
frequent deformity of the head is a marked constriction at the
posterior part so that it is the shape of a pear or top as in Fig. 27.
In some, the head is otherwise normal in size. Avhile in others it is
much elongated, as in Fig. 28, or considerably atrophied. A con-
striction at the middle of the head, as in Fig. 29, is not uncommon.
Both defects are undoubtedlj'- the result of nuclear deficiency, as
the nuclenr part of the head in these cases is much diminished in
size, and stains very deeply or not at all. I have seen spermatozoa,
the heads of which were like an inverted cone, with a bulging
rounded base. Other head.s are even somewhat contorted and bent
on themselves, as shown to some extent in Fig. 26.

Lender caudal deformities, the most frequent form encountered
is a thickening of the connecting piece. This may occur as a uni-
form thickening, or as a bulging appendage. Rudimentary de-
velopment of the tail, the presence of two poorly formed tails, and
defective development of the connecting piece occur rather infre-
quently.

All these immature and defective types are. as a rule, motion-
less, and of course incapable of producing fertilization. Their
presence indicates lowered fertility of the semen. Besides these
deformities, there are sperms showing a cur\'ature of the tail at an
acute angle just posterior to the neck, — the so-called " wry neck."
Their significance is difficult to explain, but they occur frequently
in somen fixed and stained by the same routine methods used on
samples in which thev are al)sent. They probably are not the re-
3

34

suit of the methods used in fixing and staining. Some think they
are slightly immature types, or that the condition is produced by
abnormal contractions of the tail. The majority of sperms, ho^y-
ever, especially those from highly fertile bulls, do not show this
type at all.

The most common changes in the spermatozoa, are those in
which there is a separation of the head from the tail, and degenera-
tion of the head as e^'idenced by reaction to stains. The separation
of the liead from the tail always occurs at the neck, and often is
associated with degeneration or abnormalities of the head. The
separation, in the majority of cases, indicates some lowering of
vitality in the elements, altliough in many instances traumatism
produced in making smears or collecting the samples is responsible.
Various forms of abnormal staining of the head are very common.
The cell membrane, wliich is normally distinct and sharp, becomes
blurred in outline. Normally, the head takes a good differential
stain, the anterior part staining lightly, and the posterior part
somewhat deeper. The nucleus is distinct in outline and well de-
fined. The lighter " inner body " stands out in well stained speci-
mens. As the result of degeneration, the whole head may take
the stain uniformly, either slightly or much deeper than normal,
according to the degree of degeneration. The whole problem of
staining, however, depends very much upon the methods used,
and the care with which they are applied. When a good method
is obtained, it should lie adhered to, and used uniformly on all
specimens. As a rule, however, a certain amount of practice will
enable one to differentiate between the sharply outlined, clearly
staining normal forms, and those that show abnormal reactions to
the stains.

Bacteriology

A complete bacteriological study was made of the genital tracts
of fourteen normal young veal calves (six to twelve weeks old),
four mature fertile IduIIs, and sixteen mature bulls, either sterile or
impotent to some degree. Together with these, the tracts of eleven
aborted fetuses, seven calves dying from calf infections (scours or
]:)neumonia). and sixteen bulls slaughtered at an abattoir were
studied Ijacteriologically. Occasionally, studies were made of indi-
vidual seminal vesicles or testes, when these parts alone were
brouglit or sent in. The history of the abattoir animals was, of
course, quite indefinite or entirely negative. On the killing floor,
many tracts could be studied for pathological changes, but in the
bacteriological work it was difficult to care for more than two
tracts on each visit.

The results of the bacteriological examinations are given in the
appended ta])les, in which the tracts are divided into six groups.
The results in Oroup I, consisting of normal veal calves, indicate
that the genital organs of young male calves are, under normal
conditions, free from bacteria. Carpenter (9) obtained like re-
sults in examining tlie genital tracts of heifer calves.. The cul-

35

tures made from the seminal vesicles and testes of all these veal
calves were, with two exceptions, negative. Both seminal vesicles
of one tract and one of another yielded cultures of Staphylococcus
all) us.

Adult bulls of known fertility were naturally difficult to obtain,
only the four animals in Group II being available for examination.
Two of these (Nos. 1 and 2) were from the experimental herd
kept by the department, and at all times had a good breeding
history. The other two were good breeders, but were slaughtered
because of poor pedigrees. Bull 1, raised in the department herd,
had a severe attack of scours when a few weeks old, while the calf-
hood history of the other is not known, he having been purchased
after reaching sexual maturity. The cultures from the genital
organs of the former (Bull 1) were entirely negative, exce{)t those
from the left epididymis and scrotal sac, which yielded growths of
Streptococcus viridans. All tlie organs of the tract from this ani-
mal were normal, except foi- the fact that numerous strands of
connective tissue extended from the serous covering of the tail of
both epididymes to the adjacent part of the parietal layer of the
tunica. The tract of the other failed to show any organisms. The
only evidence of any abnormality was the presence of the same
connective tissue strands on the tail of the epididj-mis, as in the
first tract. The other two Ijulls gave negative cultures from all
parts.

Of the sixteen bulls in Clronp TIT. slaughtered at abattoii-s, and
in which no history was available, eight failed to show the pres-
ence of any organisms in their genitalia. Of the others, the vesicles
yielded cultures of Stapliiilococcus alhus nine times, and strepto-
cocci four times, Stapltjflococcus aJhus was i-ecovered once from
the prostate, and once fi'om Cowper's glands. The testes gave cul-
tures of staphylococci in two cases, and Bact. ahortum in one. No
observable anatomical changes accompanied the presence of the
Rang bacillus in this case. The epididymes showed growths of
staphylococci fi^'e times, and streptococci on three occasions.
Streptococci were isolated fi"om the scrotal sacs of eight testes.

The results in Group IV (aborted fetuses) show that bacteria
are often present in the seminal vesicles or testes of these animals.
As a rule, however, the organisms are identical with those isolated
from the l)lood or other jiarts of tlie animal. This is to be ex-
pected, however, for liecause of the feeble resistance of the fetus
to any infection, the organisms circulating in the bloo<l may be
isolated, as a rnle. from many ditferent organs and tissues. All
samples of blood set with Bact. ahortum antigen were negative,
irrespective of whether or not the organism was recovered from
the blood or other tissue^. This is in accordance with the findings
of Garpenter in the female fetus. — the resistance is so feeble that
few or no antibodies are formed to combat any existing infection.
Bact. ahortum was recovered in two cases from the vesicles, and in
four cases from the testes, but in each instance the same organism
was present in the blood or otjier tissues of the body.

36

The results from the tracts of the calves dying of calf infec-
tions are given in Group V, and show tliat five were negative. The
other two showed B. coli, staphylococci, and streptococci, in the
organs indicated by the chart.

In Group VI, the mature infertile or sterile bulls, there was a
comparatively wide variation in the type of organisms encoun-
tered, but streptococci and micrococci were the most common in-
vaders. In the order of the frequency of infection, the organs
would be enumerated as follows: Vesicles, epididymis (usually
the tail), scrotal sac, testes, prostate, and Cowper's glands. The
first three parts mentioned usually contained bacteria. A strep-
tococcus Avas the usual invader of the scrotal sac, and very prob-
abl}^ was the cause of the connective tissue tufts and strands so
frequently seen. The vesicles and epididymes gave, in the order
of the frequency of their occurrence, staphylococci, streptococci,
B. coll, and Ps. pyocijaneus. The streptococci were usually of
the viridans group, though a few were hemolytic, and two strains
were indifferent to blood. The testes gave growths in only eleven
instances, — staphylococci eight times, streptococci two times, and
an unidentified rod once. The prostate yielded staphylococci twice
and Cowper's gland once.

As emphasized previously, the vesicles and tail of the epididymis
are most subject to infection and degenerative changes. At the
same time, they are intimately connected with the secretion of
the semen. Once the epididymis becomes infected, there is nothing
to prevent the organisms, together with inflammatory products,
from being mixed with the semen and ejaculated during coitus.
Also in the vesicles, unless the inflammation is so severe as to
occlude the excretory duct, the organisms are mixed with the
vesicular secretion, which is emptied into the urethra during
ejaculation. Carried along v/ith the bacteria, are, of course, toxic
products, degenerated cells, and the otherwise altered secretion of
the glands. One interesting case noted was that of a bull that
had passed from a state of fertility to that of complete sterility
during a period of two months. The semen was semi-fluid, green-
ish yellow in color, and contained a very few non-motile sperma-
tozoa. Post mortem examination showed that the vesicles had
undergone abscess formation and that they contained yellowish
green material similar to that which had been discharged during
copulation. Streptococcus hemolytic us and Ps. jyjfocyaneus were
isolated from both vesicles, and from the semen. Micrococcus alius
was isolated in nearly all cases of vesiculitis and was often asso-
ciated with Streptococcus viridaus or hemolyticus.

Bacteriological studies of the semen are, on the Avhole, more or
less unsatisfactory, due to the present difficulty in obtaining sam-
ples free from any chance of contamination. In most of the ab-
normal bulls, bacteria of various types were isloated from the
semen, most of which agreed culturally with those later isolated
from the internal genital organs of the same tracts. The method
of culturing consisted of douching the prepuce of the bull and

37

vagina of the female with sterile saline solution before breeding.
Samples of vaginal mucus were taken before service, and the flora
compared to that after douching. This method of douching pro-
duced vaginal samples relatively free from bacteria, at least so
much so that the post coital fluid demonstrated that many organ-
isms must have been introduced from without. AA^hether or not
they came in with the semen is problematical, but in all probability
this was the method of introduction.

I have so far failed to obtain Bad. ahortum from the tract of an
adult animal, either by direct culture or guinea pig inoculation,
except from the testicle of one abattoir bull. The agglutination
tests with Bad. ahortum antigen were all negative, except for tAvo
abattoir bulls. The results so far obtained would seem to indicate
that, in accordance with the findings of other workers, the Bang
organism seldom invades the male genital tract, or does not thrive
there after its introduction. Schroeder (12) and others, have,
however, on various occasions, recovered the organism from the
bull, and the former author even states that it invades the vesicles
and is eliminated with the semen.

Discussion

A complete discussion of those factors which have a bearing on
reproduction and fertility in an animal, includes not only a
thorough study of the genital tract, but an appreciative considera-
tion of various extrinsic factors. The effect of environment has
long been known to have a marked influence upon breeding, par-
ticularly Avith reference to animals in domestication. Diet, though
long relegated to a minor phase of the question, has, Avithin recent
years, come to be a matter of prime importance AA'ith regard to its
bearing upon the entire body metabolism. The endocrine organs
preside over and regulate the gi'owth and functioning of the geni-
tal organs from the earliest embryonic stage to the cessation of
sexual life. Any derangement in one results in functional or
organic changes in the other. In a given mating, aa^c must take
into consideration such factors as impediments to coitus, as well
as those numerous agencies in the female AA^hieh may interfere with
the union of sperm and OAaim, or AA'ith the successful implantation
of the fertilized eg^^ in the uterus, and its groAvth and deA^elop-
ment there till normal parturition takes place. Successful repro-
duction depends upon the mating of sexually sound females to
equally sound males. Considering the A^arious factors AA-hich gov-
ern reproduction, sexual soundness must necessarily depend, to
a large extent, upon a good general condition of the entire body.

ExA'iRONMENT: The effeet of environment on fertility in the
bull is no doubt a minor factor. Cases in Avhich changes in en-
vironment affect fertility probal)ly occur, liOAvcA-er, particularly
when fear and other psychic disturbances play a part. Marshill
(29) states: "Tt Avould seem probable that failure to breed
among animals in a strange enA'ironment is due not, as has been

38

sxinrtrested, to any toxic influence on the organs of generation, but
to the same causes as tho.se which restrict breeding in a state of
nature to certain particular seasons, and that the sexual instinct
can only be called into play in response to certain stimuli, — the
existence of which depends to a large extent upon appropriate
seasonal and climatic changes."

Diet: Under tliis heading we may include not only the effect
of deficient food, but also constitutional disorders, as a result of
"which the oi'gans of generation and those glands guarding their
function I'cceive insufticient nourishment. It is a well known fact,
and long has l>cen. that animals fail to breed when they are in a
run down condition or when they are fed a deficient diet. Gary
(35), quoting Hagner, states that the virility of the spermatozoa
is often in direct proportion to the general physical condition of
the patient.

Reynolds (di) emphasizes the fact that it is an established
l^rinciple among animal breeders that a high protein diet in both
sexes is essential to full fertility. " Oligospermia with defieient
vitality of the spermatozoa is not infrequently found from con-
stitutional disorders. It can easily be demonstrated in animals
that both low diet and conditions of life that produce a nervous
excitable state are attended by oligospermia." Animals that are
closely confined, those that are over-fat (show animals), as well
as those fed a deficient ration very frequently fail to breed,' but
exercise and change of diet soon overcome the impotency.

Dutscher, Hart, Steenbock. and other biological chemists have
done extensive work to show the essential importance of vitamines
and minerals in the diet. Their results indicate that animals can-
not thrive and breed normally when fed a diet composed solely of
the products of one plant. There must be variety, and there must
be not only a correct nutritive ratio, but the mineral and vitamine
content must be present as well. Cows fed on the products of one
plant often failed to ])reed, and if conception occurred, it in-
variably^ resulted in a premature birth, or the birth of weak and
poorly nourished calves. The work of these authors is funda-
mental, and Ijrings out many important points. Is it not probable
that the deficient diet results in Aveakened tissues which are easier
proy to the invasion of bacteria?

Macomber and Reynolds (39) experimented upon white rats
to determine the effect of defective diet as a cause of .sterility.
They call attention to the confusion caused by the application of
the term sterility to most, or all, infertile matings. They believe
that failure of reproduction is, in fact, the result of cleereased
fertility rather than of actual sterility on the part of the two indi-
viduals concerned. " There are certainly a large number of in-
fertile matings which are purely functional and due to physiologic
alterations or local conditions. Such physiological alterations more-
over coexist in the sterilities of ])atholoo:ic origin and when un-
recognized and consequently unremedied, undoubtedly explain a

39

large proportion of tlu' continued infertilities after operation."
Is it possible for a bull to be infertile to the cows in his herd
that have been fed a deficient diet, and at the same time to breed
well when mated to animals outside this herd ? This is rather im-
probable in practice, but there is always the possibility of its
oceurrenee. In tlie experimental work, white rats were used: one
strain from tlie \Vistar Institute with a fertility of about 65 per
cent, and the other from a Dr. Castle's strain with a fertility of
about 90 per cent. The "Wistar rats were fed in groups, each
group receiving a diet deficient in a certain sul)stance : calcium,
protein, or fat soluble vitamine. To this grou]) was added a diet
deficient in both calcium and protein (war diet). These diets
reduced the fertility of the groups from the original 65 per cent,
to 55, ?)1. and 14 per cent respectively. It delayed the appearance
of fertilitv in voung rats raised on these diets, and lowered its
degree in the mature animals. Most of these rats, however, though
infertile to each other, bred promptly Avhen mated to the Castle
rats of known fertility. This demonstrates clearly that relative
infertility of given matings does occur. One interesting feature
of the work is the fact that in the matings on the single deficiency
diets, four deliveries of macerated fetuses occurred and there were
two more in eight deliveries from those reared on the war diet.
Xo cases of this kind had previously occurred in this strain, which
had 1)ecn under observation for several years. Does this throw
any light upon the cause of macerated fetuses in cattle? Micro-
sco]iically the testes and ovaries of these infertile rats showed no
observable changes, a fact which is of great importance to bear in
mind.

Williams, in his book on disease of the genital organs, brings out
quite clearly the relation of defective diet, overfeeding, and lack
of exercise, to reproductive efficiency.

Xovarro (40) observed that pigeons fed on a diet without vita-
mine B showed degeneration of the seminal epithelium, with
hypertrophy and hyperplasia of the interstitial cells of the testis.
Another author (Allen) .showed that reduction in the quantity of
water-soluble vitamine in the diet of rats resulted in total degenera-
tion of all the germ cells, but it did not interfere witli gi'owth and
development in other respects.

The observations of Williams (41), in a pure bred beef herd in
Hawaii, clearly demonstrate the intimate correlation between poor
fodder as the result of extreme drought, and the accentuation of.
or increased susceptibility to, genital infections, as demonstrated
by clinical findings. The genital disorders started soon after the
onset of the drought, and immediately took a downward trend
with the advent of the rainy season.

Judging by the work quoted, we will observe tliat deficient diet,
though it does not always affect the general health, has a ]irofnund
effect upon the genital organs of l)oth sexes, associated with dis-
turbances of spermatogenesis in the male. In most debilitated and

40

weakened conditions of the male, spermatogenesis ceases or is
markedly defective. We must, undoubtedly, explain this fact
upon the ground of deficient nouHshment to the reproductive
organ.s or possibly the endocrines. The vitamines have been
termed nuclear nourishers, and their absence probably results in
nuclear deficienc}'.

ExDOCRiNES: Bell (42) emphasizes the fact that not only the
structure but also the function of every part of the body is in
close correlation with the rest. " This is essentially true of the
ductless glands: the shadow of their influence is over all."
Further he states that when we remember that the individual
exists to perpetuate the species, it is not difficult to realize that
the metabolic factors concerned in reproduction are the same as
those related to the individual metabolism. It follows, therefore,
that the ductless glands which regulate the individual metabolism
concern equally the reproductive. Brown (43), discussing the
same subject brings out the generalization that the sympathetic,
since it is the most primitive part of the nervous system, is closely
associated with the endocrine system, a still more elemental means
of communication in the body. Also since specialized reproductive
cells appear before the nervous system, the organs of reproduction
remain closely associated with the older chemical reactions now
specialized in the endocrine glands. " The endocrine glands, the
reproductive organs, and the sympathetic nervous system, there-
fore, remain as a basic tripod, and it is not likely that a disturb-
ance will occur for long in one limb without affecting the other
two." The former author believes that the gonad keeps the other
ductless glands informed of the needs of the genital tract, they in
turn influencing the general metabolism. Jump (44) states:
" We are therefore justified in believing that there is a correlation
of function between these (endocrine) glands and that some cases
of sterility are probably due to a derangement of this correlation."
Biedl (45 ") concludes: " There appears to be an intimate anatomi-
cal and physiological interrelationship between the different blood
glands which is manifested clinically by the fact that the patho-
logical disturl)ance of one gland is accompanied by symptoms point-
ins: to the functional derangement of one or more of the others.
Knowing, as we do, the many sided interactivity which subsists be-
tween the different internal secretory organs, itTs readily con-
ceivable that isolated diseases of single organs of this group are
very much rarer than, at the first glance, they would appear to be.
Tn the present state of our knowledge, the only course of investiga-
tion which is open to us is to start with the known results of the
functional derangement of any organ, and, by following these up,
to seek the primary link in the pathological chain."

Most workers seom to agree that a special connection exists be-
tween the normal function of the adrenal cortex and the sexual
organs. Tumors of the former are usually associated with sex

41

abnormalities, and feeding young animals the gland substance
seems to stimulate growth or the testes.

Many arguments have been brought forward to show that the
prostate produces an internal secretion. It is a well known fact
that this organ atrophies after castration, and enlarges as the sex
life dwindles. As has been previously stated, Serrlach and Pares
reached the conclusion that the gland prodvices an internal secre-
tion which controls the testicular functions and regulates the pro-
cess of ejaculation. Also they state that if the prostate is re-
moved, spermatozoa are no longer produced in the testes, and that
the secretory activity of the accessory genital gland ceases. The
secretion is, at any rate, a stimulus to the internal secretion of the
testis.

The thyroid bears a distinct biological relationship to the sexual
glands. Removal of the gland results in imperfect development of
the gondas. infantilism, and general torpor. Bell (42) believes
that the association between ^:he thyroid gland and the genitalia is
as intimate as the relation of the pituitary to the genital functions.

Of all the endocrines, perhaps the anterior lobe of the hypophysis
is in most intimate correlation with reproduction. Castration
results in hypertrophy of this organ, while removal of the anterior
lobe usually leads to death. In those cases in which death does
not ensue, it results in genital atrophy, stunting, and- reduc-
tion of sexual activity. In young animals, spermatogenesis ceases
entirely even after partial extirpation of the anterior lobe, Biedl
(45) states that " in disease of the hypophysis, derangement of
sexual acti^-ity occurs very early in the course of the disease. sho'v\Ti
in women by the cessation of menstruation, and in men by im-
potence,"

The thymus, as is well known, is quite intimately associated
with the development of the genital organs. Its normal disap-
pearance is always associated with the development of sexual
maturity in the individual. Hewer (46) conducted experiments to
ascertain the effect of thymus feeding on the activity of the re-
productive organs in the rat. She concludes in part : ' ' Male rats
appear more susceptible to the influence of thymus feeding than
female rats. With moderate doses of thymus, sexual maturity in
the animals treated is delayed, a phenomenon which is attributed
to delayed development of the testis. AYith large doses of th\nnus,
in the male, the testis is structurally affected : in the young animal
in the direction of retardation of development, in the mature ani-
mal in the direction of degeneration. This degeneration is con-
fined to the testes. In the degenerating testis, cells of Sertoli ap-
pear to be absent : the spermatogonia are present, also dividing, and
may lie free in the lumen of the tubule ; spermatids, many with
abnormal nuclei, are shed into the lumen in large numbers ; sper-
matozoa are practically nbsent. In the later stages, only a few
dividing spermatogonia appear among the debris of the other un-
recognizable cells of the tubide. In the epididymis Avhich itself is

42

normal, when the testis is showing degeneration, ver^" few spei*ma-
tozoa appear, in the later stages none. Many spermatids are pres-
ent in various stages, and some spermatocytes. Animals in the
hyper-thymic condition appear to be sterile."

The foregoing references will, I hope, serve to bring out the facts
that environment and diet, together Avith the general body meta-
bolism and the endocrines, have a more or less profound effect
upon the development, growth, and functioning of the genital
system. In the experiments it has been shown that sterility is
not necessarily accompanied by any apparent microscopical
changes in the gonads, or even at times in the general body health.
Nor can w^e exclude impotency of the male entirely even when the
spermatozoa are normal in shape, and motility. Carnett, and
others (38), years ago stated: " Indeed, there is abundant clini-
cal proof to the effect that systemic conditions which have no ap-
preciable effect upon the motility or conformation of the sperma-
tozoa materially interfere with reproductive power." The entire
complex genital system is inseparably linked up with the body as a
whole, a fact which we must bear in mind at all times.

Impediments to coitus may be due to great difference in the size
of the two mated individuals, psychic disturbances, or inability
to protrude the penis. Williams (17) mentions several physical
impediments, as deformity of the limbs or feet, sore feet, overload-
ing of the rumen, obesity, fear of falling, and paralysis. Coitus
may be somewhat delayed, or even not performed as the result of
a severe inflammation with sensitiveness of the penis or prepuce.
Occasionally tumors of the penis are encountered which may inter-
fere with prolusion of the penis, or its entrance into the vagina.
Not infrequently the penis is rendered incapable of protrusion as
the result of inflammatory adhesions, tuberculosis of the preputial
lymph glands, etc.

Excessive sexual use, within certain limits, probably has not, in
itself, any material permanent effect upon the reproductive capac-
it}-. Tlie frequency with which bulls used to excess break down
sexually, is probably due to the devitalizing effect upon the tissues
of tlie genital organs, this opening the way to bacterial invasion
and other destructive influences. Over-use is probably not danger-
ous, unless continued over long ]ieriods. but at the same time it
offers greater opportunity for infection to be introduced into the
body from intercourse with large numbers of females. Lloyd-
Jones and Hays (47) carried on very interesting experiments on
the influence of excessive sexual activity of male rabbits on the
properties of the semen. Their plan was to mate male rabbits in
quick succession, and study the character of the semen on the
first service, and every fifth service thereafter. The safe limit
was twenty services in three hours. As would be expected, the
volume of the semen, after the first few services, became gradually
reduced in amount. " In rapidly successive services, the semen
becomes less viscous and tends to lose its charactertistic milky

43

appearance until at the twentieth service, when the fluid is thin
and watery," It seemed as though there was a well marked reduc-
tion in the number of spermatozoa per cubic centimeter in the ad-
vanced servdces. Successive copulations also resulted in a marked
decrease in the number of motile spermatozoa, together with a
sliorter duration of perceptible vitality. The certainty of produc-
ing impregnation at the same time became less and less. "This
reduction in the per cent of effective matings when the male is
sexually overworked is recognized hy those engaged in animal
breeding as one of the most noticeable and universal concomitants
of heavy sexual service."

In another paper, these same authors studied the effect of sexual
excess upon the character of the offspring. In part, they conclude :
''Ba' no means thus far used has any inferiority of i)rogeny from
the heavy sexual service been discovered. They are fully equal if
not superior to progeny from very light service of male."

Infection is without doubt the greatest single factor capable of
producing functional and anatomic changes resulting in varying
degrees of imi)oteiicy and sterility. The changes produced range
from the addition of the toxic products of bacterial growth to the
seminal fluid, to the complete destruction of the parenchymatous
tissue of one or more of the contributing sextial glands. Anatomic
changes are l)y no means essential to the ])roduction of lowered
fertility. As has been previously stated, the work on veal calves
indicates that the genital organs of young bulls are normally free
from bacteria. Likewise in normal adult animals, the bacterial
content of the genital organs is as a rule low or negative. It is
possil)le that a certain flora is normal for the tract at sexual
maturity, as in several other organs of the body, but under the
strain of sexual excess, defective diet, or other weakening in-
fluences, the5-e organisms may become pathogenic. Streptococci and
staphylococci have at times been found in apparently normal parts
of the body, and at other times they are found associated with
severe pathological lesions in the genital tract. The degree of
pathogenicity is of course difficult to determine, except as we find
the bacteria associated with abnormal conditions. Carpenter (9),
however, injected streptococci into the genital tracts of female
calves and produced lesions resem])ling very closely those from
which the organisms had been isolated in adult sterile animals. Per-
sonally. I am inclined to believe that the genital organs normally
are free from bacteria, or if any are there they are better able
to multiply under the strain of devitalization of the tis.sues.
Bacterial invasion, liOAvever, does take ]ilace fpiito fi-equently. l)ut
the paths of entrance of the organisms are .somewhat proble-
matical. Hematogenous origin is always possible, though it is
rather difficult to definitely implicate this mode of entrance. The
urethra is perhaps the easiest and most common ]iath for the
entrance of bacteria, though even here it is not ])Ossible to make
definite assertions. Contiguous spread of infection from neigh-

44

boring structures is very probable iu some cases, particularly in
pelvic peritonitis. The Ijaeteriological results hardly bear out the
theory of ^Yilliams that the organisms lie dormant in the genitalia
of the animals until the advent of sexual maturitj^, at which time
they acquire pathogenic powers. On the other hand, his clinical
observations seem to indicate that this may be possible. Calves
suffering from "calf infections" frequently do harbor organisms
in their genital organs, but whether or not they persist there till
sexual maturity is a matter of conjecture. The most logical theory
seems to be that animals from herds in which genital infections
are very severe, or those that have had severe attacl<s of scours
or pneumonia, are more susceptible to those infections, due to the
early lowering of their vitality. One bull in the department herd
certainly had a severe ordeal as a calf, but as a mature bull he
was highly fertile. Moderate sexual use and proper sexual hygiene
proljably had much to do with this. In the bull, infectjon of
some part of the genitals, during some period of life, is very con-
stant, however, whether or not it is productive of observable
changes in his breeding efficiency. The finding of the fine connec-
tive tissue strands and tufts on the serous surface of the tail of
the epididymis of practically all bulls examined, both sterile and
fertile, indicates past or present infection of the scrotal sac. The
vesicles and tail of the epididymis are, as stated previously, the
most commonly invaded tissues of the tract. The testes are less
frequently involved.

While it is difficult to obtain irreproachable proof that the bull
is a disseminator of genital infections, the findings of clinicians
quite clearly indicate that this is true, and laboratory methods
tend to support this assumption. Williams believes that not only
may the bull infect the female with organisms whicli interfere
with the given conception, but that he often implants there organ-
isms which interfere with future pregnancies, and even with the
life of the individual in some cases. The high abortion and ster-
ility rate following the use of certain sires, and the appearance of
characteristic infections after seiwice to certain bulls, clearlj^ in-
dicate that in all probability the bull does eliminate with his
semen those organisms whicli produce lesions in his genital organs,
and are capable of infecting the female. W. L. Williams (48)
cites the case of a pure bred herd in which breeding had pro-
gressed satisfactorily until heifers had grown to breeding age and
a second bull was obtained. " Some cows of the old herd were
also assigned to the young bull which had not previously been in
service. The cows bred to the old herd bull continued to breed
normally. The cows and heifers bred to the new bull conceived
with difficulty or not at all. Those which conceived mostly
aborted, and those which calved had metritis and retained fetal
membranes. The two first cows in which pregnancy terminated
died of metritis." I have frequently had semen samples sent in
from bulls that were not only failing to get cows with calf, but

45

following- each service the females showed a severe vaginitis.
\V. W. ^\ illianis worked in a herd in which service to certain bulls
was in each case followed by a severe vaginitis and cervicitis, only
to be followed later by a characteristic salpingitis.

Vaginal smears taken before and after service, in many in-
stances, show that in all probability bacteria, especially the strep-
tococci, were deposited there with the semen. These results have
been obtained upon several occasions, at which time the vagina
was usually douched prior to each service with sterile saline solu-
tion. Streptococci and other organisms have been isolated from
the vaginal samples obtained by this method. In most cases, they
were absent from samples taken before service. Extraneous con-
tamination, and error, must l^e taken into consideration, but the
results tend to bear out clinical observations that the bull is prob-
ably a disseminator of some infections associated with the genital
organs of both sexes. At any rate, organisms have been isolated
repeatedly from the genital organs of the bull, of the same bio-
logical character as those which are associated with sterility, abor-
tion, and allied phenomena in the female. In the absence of
obstruction in any part of the tract, there is nothing to hinder
infection from gaining access to the seminal fluid, and being ex-
creted during ejaculation.

Hopper (18) states: "A diseased bull may manifest non-fertility
or decreased potency in different ways — by repeated service to
apparently normal females without conception, by a high abortion
rate in females that have been apparently normal, by characteristic
infections following the use of any particular sire, or by abnor-
malities in the breeding tract noted by rectal or physical pal-
pation."

Admittedly, Bad. abortum has little affinity for the genitalia
of the bull, though Sehroeder states that the bull harbors the
organisms in his seminal vesicles and that they are eliminated
with the semen. Other authors have occasionally isolated the or-
ganism from the vesicles, testes, or both. Sehroeder 's theory that
infection of the female occurs indirectly by contamination of the
fo<lder with the semen is prol)ably rare in occurrence. The very
limited number of cases in which investigators have demonstrated
the presence of the organism in the male genital organs, and the
apparent immunity of the bull to the bacterium as determined by
the agglutination reaction, seem to indicate that he plays a small
part in the spread of this type of infection in the here!. On the
other hand, it would seem that he is at times intimately associated
with the spread of certain other organisms that interfere seriously
with herd reproductivity.

The diagnosis of infertility and sterility rests upon a thorough
physical examination of the genital organs, together with a de-
tailed study of the semen. The histoiy of the animal and herd
involved must also be verj'- carefully inquired into, especially the
part covering the result of everj' serv'ice by the sire in question.

46

Besides this, we must always consider all factors which have a
bearing upon the subject, remembering the phj'siology of each
part, and the role it plays in reproduction. Bacteria gain entrance
to many parts of the tract, where they multiply and probably add
toxic products to the seminal fluid, altering its biological char-
acter and resulting in partial or total destruction of the secretory
tissues. The testes, epididymes, vesicles, prostate, and other parts,
each contribute their essential part to the semen, abnormalities of
any one of which, as a rule, result in interferences with reproduc-
tion. If the vesicles are involved, we must bear in mind just what
is the part played by their secretion, and what is the probable
result if their essential elements are not added to the semen. In
like manner, we must consider the prostate, whose secretion stimu-
lates the vitality of the sperms, and adds fluid bulk to the semen.
Extirpation of the vesicles or prostate alone results in lowered
fertility, without altering the sexual desire, while removing lioth
glands produces total sterility. Partial or total destruction of the
parenchyma of either gland produces the same effect as extirpa-
tion, in that its function is altered or entirely absent.

The semen should be examined, not only for the number of
spermatozoa and the percentage of those that are motile, but for
the duration and type of motion. Normal semen, when first ex-
amined under the microscope, shows a field closely packed with
highly motile spermatozoa. In every study of the semen, however,
we must 1)ear in mind the temperature and other conditions under
which it has been kept since emission. On the other hand, semen
from bulls of lowered fertility shows changes ranging from mild
disturbances such as sluggish motility and a slight decrease in the
number of sperms present, to aspermia, or total lack of motion.
Normal semen, Avhen compared with abnormal specimens, as a rule
presents distinct differences, either in motility, staining properties,
or structure of the spermatozoa. Impotent bulls, however, may
show at times few or no ol)servable changes in their genital organs.
The only assumption hei-e is that the condition probably is of
endocrine origin, or is some functional disturbance. Of oligo-,
spermia Reynolds states: ''Oligospermia, Avith normal motility
and vitality, is not al)so]ute sterility, l)ut is of high importance
because the percentage of destruction of spermatozoa during their
passage through the genital canal of the female is so enormous that
the possibility of impregnation by semen which starts out with a
deficient number is always poor. When the genitals of the female
partner are in a condition which is even moderately hostile to the
spermatozoa, impregnation by such semen becomes so unlikely as
to be not even a probability."

Motility may be lacking in a small number, its absence may be
observed in a large percentage, or even in all those in the field.
as in necrospermia. On the other hand, the motility may be
sluggish or of abnormal ty])es in variable percentages. Sperms

47

with sluggish motility are always low in vitality, and have weak
powers of insemination, as the motion lasts but a comparatively
short time. The vitality may be but moderately lowered, so that
although the sperms are highly motile when ejaculated they soon
lose their power of propulsion. The type of motion is likewise an
indicator of lack of vitality. The '"progressive vibratilc" motion
described by Reynolds should proceed to a high degree for a long
period before the "undulatory tactile" or bunting types of motion
appear. Early appearance of these two latter ty])es indicates in
most eases a marked lack of vitality of the elements. The motion
should be vigorous and lasting, for, as stated by Reynolds, "noth-
ing is more certain than that spermatozjoa of merely moderate
vitality seldom impregnate a female."

The early precipitation out of the "Boettcherchen" crystals is
very characteristic of oligospermia, and impotent semen. The
theory here is that crystals do not precipitate out wdien a fluid i-^
actively moving, but soon do so when the fluid is motionless. The
sediment which normally makes up about two-thirds of the sample
is usually decreased in abnormal samples. The semen itself should
be observed for unusual viscosity or a thin watery condition.
Clotting should occur readily after emission, but the clot soon
liquefies to some extent, allow'ing the spermatozoa to become more
active. This clotting is, of course, to protect the delicate spei-ms
from the hostile secretions of the vagina.

The presence of immature and deformed types of sperms rep-
resents some disturbance of spermatogenesis, but it is difficvilt to
explain the significance of these forms. They are seldom seen in
normal samples, and undoubtedly none are capable of producing
impregnation. In the case of minor abnormalities of staining rf-
actions. the sperms are probably deficient in nuclear material or
othei-wise altered so that probably they are incapable of reaching
and uniting with the ovum. When impregnation does occur in
the.se cases, weak offspring undoubtedly result in many instances.

The work has by no means progressed to the point where one
may. by an examination of the semen, determine the degree of
impotency with great accuracy, or even whether the animal may
be restored to sexual health by proper hygienic and therapeutic
treatment. Ivelatively, the greater the changes in the semen and
spermatozoa, the less the chances of impregnation. Infertility
to any marked degree, is, however, usually accompanied by cor-
responding ehanges in the seminal fluid and its germinal elements.

Examination of the semen is, and pro])ably always will be.
simply an aid in reaching a diagno.sis. AVhile abnormalities of
the semen and spermatozoa are associated Avith sterility or in-
fertility, it is unwise to lay too much emphasis upon this method
of diagnosis alone, especially Avith regard to the making of a
definite prognosis. When large numbers of abnormal spermatozoa
are present in the semen, we are safe in saying that the animaJ

48

is, at the time, of lowered degree of fertility. One should be very
cautious, however, in foretelling how long the condition will last,
or if the animal may in time be restored to full fertility. Ster-
ility, due to' organic disturbances, probably seldom yields to treat-
ment, but when it is due to functional disorders resulting from
defective diet or lack of exercise, the condition is frequently reme-
died by overcoming the cause. Lack of exercise and overfeeding
seem to be etiological factors in a fair percentage of cases.

Besides abnormalities of the male genital tract, we must always
consider the numerous factors in the female that may kill or
weaken the sperms. Impediments to successful coitus may be
present in the form of vaginal constrictions, abnonnally short or
small vagina, or other deformities. Hostile exudates, mechanical
obstructions, and other factors may interrupt the progress of the
sperms at any point in the tract.

Although little is known definitely regarding disorders of the
endocrines in the bull and their relation to reproduction, the w^ork
in human medicine and experimental researches upon laboratory
animals warrant thoughtful consideration of these factors which
are by no means insignificant. In the future, these glands will no
doubt receive more and more attention in their relation to the
genital organs and reproduction.

Conclusions

1. The genital organs of the bull quite frequently undergo
pathological changes, due to infection with the same varieties of
microorganisms associated with genital infections in the female.

2. In all probability, these microorganisms are frequently elimi-
nated with the semen and infect the female during copulation.

3. Past or present infection in the genital organs of all the
bulls so far examined was evidenced by the presence of the fine
connective tissue tufts and strands upon the tunica vaginalis, par-
ticularly that part covering the tail of the epididymis.

4. LoAvered sexual capacity is, as a rule, accompanied by
demonstrable changes in the semen.

5. A study of impotency and sterility includes not only a thor-
ough study "of the genital organs, but also those extrinsic factors
which govern reproduction either directly or indirectly.

6. A thorough knowledge of the anatomy and physiology of the
male genital organs is fundamental to a clear understanding of
the problem.

I am much imlebtecl to Drs. W. L. and W. W. Williams for some of the
material, and for helpful co-operation in the early part of the work; to
Drs. C. M. Carpenter and R. R. Birch for many helpful suggestions; and to
Dr. J. N. Frost and others who so kindly co-operated by placing at my dis-
posal samples of semen and some of the genital tracts.

49

Bacteriology of the Genital Tracts of Normal Young Cal\^s

Group I

Number

Right sem. vesicle Left sem. vesicle

1

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Left testia

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Bibliography

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Med. Band I, S. 241, 1897. Das seucheiiliaften Verwerfen der Riiidoi'.
Arch. wiss. u. prakt. Thierlieilk., Band 33, S. 312, 1007.

2. Law, James. Contagious Abortion of Cows. Circular No. 5, Ayr.

Experiment Sta. University of California, June, ino.S.

3. Report of Engli.sli Commission of Epizootic Abortion. Appendix to Part

I. p. 17, 1909.

4. Hadlky, F. B.. and Lotue, H. The Bull a? a Disseminator of Contagious

Abortion. Jour. Amer. Vet. Med. Assoc, L, 1916-17, p. 143.

5. Hadley, F. B. Contagious Abortion Questions Answered. Bulletin No.

296, Oct. 1921, Agr. Experiment Sta., University of Wisconsin.

6. Buck, J. M., Creech, G. T., and Ladson, H. H. Bacterium Abortus Infec-

tion of Bulls (Preliminary Report). Jour. Agr. Research, August,
1919.

7. Schrofjjer, E. C, and Cotto.v, \V. E. The Bull as a Factor in Abortion

Disease.
S. Cotton. W. E. Proceedings of A. V. M. A., p. 851, 1913.
9. Carpenter, C. M. Report of the New York State Veterinary College,

Cornell University, 1920-21.

10. Rettger, L. F., and White, G. C. Infectious Abortion in Cattle. Storrs

Agr. Exp. Sta. Bulletin No. 93. January, 1916.

11. McFadyean, Sheather and Minp:tt. Researches Regarding Epizootic

Abortion. Jour, of Comp. Path, and Therap., XXVI, 142, 191.3.

12. Sciiroedek. E. C. Bureau of Animal Industry Investigations of Bovine

Infections Abortion. Jour. Amer. Vet. Med. Assoc, LX, p. 542.
February, 1922.

13. Barney, J. D. Recent Studies on the Patliology of the Seminal Vesicles.

Bost. Med. and Surg. Jour.. CLXXI, 1914, 59.

14. WiLLi.\MS, W. L. The Diseases of Bulls. Cornell Veterinarian. X, 94.

April, 1920.
J 5. Williams, W. L. Report of the New York State Veterinary College,
Cornell University, 1920-21.

16. Williams, W. W. Technique of Collecting Semen for LalK)ratory

Examination with Review of Several Diseased Bulls. Cornell Vet.
X, 87, April, 1920.

17. Williams, W. W. Diseases of the Bull Interfering With Reproduction.

Jour, of Amer. Vet. Med. Assoc, LVIII. 29, October, 1920.

18. Hopper, E. B. Herd EfTiciencv from the Standpoint of the Veterinarian.

North Amer. Vet., III. 71, February, 1922.

19. Williams, W. W. Observations on Reproduction in a Purebred Dairy

Herd. Cornell Veterinarian, XII, 19, January, 1922.

20. Walker. K. M. The Diagnosis and Treatment of Sterilitv in tlie ^tale.

Lancet, CCI, 228. July 30, 1921.

21. Pende. Endocrinoglia-Pathologia E Clinica. Abstract in Endocrinology,

II, 42.

22. Ferguson, J. S. Normal Histology and Microscopical Anatomy.

23. Ellenberger. Vergl. mikroskop. Anatomie der Ilaustiere. II.

24. CouRRiER, M. R. On tlie Existence of a Secretion of the Epididymis of

the Hibernating Bat and Its Significance. C. R. Soc. de Biol. (Paris),
1920. LXXXIII, 67-69.
23. Stigler, R. Abstract in Jour. Phys. et Path. Gen. (2) CLXXI, 273.
September, 1918.

26. Disseliiorst. Ansfuhrapparat und Anhangsdrusen der mannlichen

Geschlechtsorgane. Oppel's Lehrbucli der vergleichenden mikr.
Anat. der Wirlieltiere, IV, Jena, 1904.

27. Rytina, a. a. The \'erumontanum. With Special Reference to the Sinus

Pocularis: Its Histology', Anatomy, and Plivsiology. Jour. Urology.
I, 1917, 231.

28. Fish, P. A. The Spermatic Secretion and Its Ultrai«rticles. Cornell

Veterinarian, October, 1921.

60

29. Marshall, F. H. A. The Physiology of Reproduction.

30. Kingsbury, B. F. Professor of Histology and Embryology, Cornell Uni-

versity, Ithaca, New York.
31. Walker. Arch. f. Anat. u. Entwicklungsgesch., 1899, iind Arch. f. Anat.
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32. BoETTCHER, W. On the Significance of the Secretions of the Male Acces-

sory Genital Organs. Miinch. med. Wchnschr. 1920, 67, 1, p. 44.

33. Editorial. Jour, of Amer. Med. Assoc, LXXVII, No. 1, 42, July 2, 1921.

34. Reynolds, E. Fertility and Sterility. Jour, of Amer. Med. Assoc,

LXVII, 1193-11199, October 21, 1916.

35. Gary, W. H. Examination of Semen With Special Reference to Its

Gynecological Aspects. Amer. Jour, of Obstetrics and Diseases of
Women and Children. LXXIV, No. 4, 1916.

36. Wolf, C. G. L. The Survival of Motility in Mammalian Spermatozoa.

Jour, of Physiol. IV, 246, August 3, 1921.

37. Brown, J. H. The Use of Blood Agar for the Study of Streptococci.

Monograph of the Rockefeller Institute for Med. Res., 1919, IX.

38. Garnett, J. B., and others. The Surgical Treatment of Sterility due to

Obstruction at the Epididymis Together with a Study^ of the
Morphology of Human Spermatozoa. Univ. of Penn. Med. Bulletin,
March, 1902.

39. Reynolds, E. and ]VL\(X>mber. D. Defective Diet as a Cause of Sterility.

Jour, of Amer. Med. Assoc, LXXVII, 169, July 16, 1921.

40. NovARRO, p. A. Tissues of the Testicle and Antaminosis. Gazetta degli

Ospedali (Milano), 1920, XLI, 424.

41. Williams, W. L. Observations on Reproduction in a Pure Bred Beef

Herd. Cornell Veterinarian. January, 1922.

42. Bell, W. B. Correlation of Function: With Special Reference to the

Organs of Internal Secretion and the Reproductive System. Brit.
Med. Jour., 1920, 1, 787.

43. Brown, W. L. The Principles of Internal Secretion. Brit. Med. Jour.

1920, II, 687-691.

44. Jump. Discussion in Penn. Med. Jour., XXV, 81.
4.5. BiEDL, A. The Internal Secretory Organs.

46. Hewer, E. E. The Effect of Tliymus Feeding on the Activity of the

Reproductive Organs of the Rat. Jour, of Physiology, XLVII, lf)13-
1914, 479.

47. Li.OYD-JoNES, 0. and Hays, F. A. The Influence of Excessive Sexual

Activity of Male Rabbits. 1. On the Properties of the Seminal Dis-
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North Amer. Veterinarian, III, May, 1922.

Description of Plates
Plate I.

Fig. 1. Diagrammatic sketches showing the development of a spermato-
zoon from a spermatogonium.

Fig. 2. Diagrammatic sketch showing the minute structure of a sper-
matozoon. The middle piece is made comparatively thick in
order to bring out the finer structures. Adapted from
EUenljerger.
Plate II.

Fig. 3. Testicle of bull, showing extensive degeneration and necrosis.

Plate III.

Fig. 4. Inner part of wall of ductus deferens. Normal, x 230.

Fig. 5. Same, Imt showing extensive degeneration and exfoliation of
the lining membrane, x 230.
Pt-Atb IV.

Fig. 6. Ductus deferens, showing entire exfoliation of the lining mem-
brane. The lumen is filled with a cellular debris, x 50.

61

Fig. 7. Same, showing the degeneration of the lining membrane, and
debris in lumen, x 230.
Plate V.

Fig. S. Seminal vesicle of bull. High power section showing the
normal structure of the vesicular cavities.

Fig. 9. Seminal vesicle of bull. Low power. The meml>rane lining the
cavities is degenerated and exfoliate<i. The cavities are
filled with cellular debris, and exudates. There is some
increase in the interstitial tissue, and atrophy of some of
vesicular cavities.
PXATE VI,

Fig. 10. Low power section of seminal vesicle of bull. The condition is
about the same as in Fig. 9 except that it is not quite as
severe.

Fig. 11. Same as Fig. 10. High power.

Plate VII.

Fig. 12. Testicular tubule showing normal spermatogenesis, x 230.
Fig. Il3. Testicular tubule showing no evidence of mitosis, x 230.
Fig. 14. Testicular tubule. The spermat-ogenic epithelium is beginning

to degenerate and become cast off into the lumen, x 230.
Fig. 15. Testicular tubule, showing almost total exfoliation of the

spermatogenic epithelium, x 230.
Plate VIII.

Fig. 16. Testicular tubule. The seminal epithelium is entirely degen-
erated. The membrana propria is markedly thickened.
X 230.

Fig. 17. Testicular tubule. The tubule is undergoing atrophy and
degeneration. The interstitial connective tissue is much
increased in amount, x 230.

Fig. 18. Same as Fig. 17, except that it is in the more advanced stages.

Fig. 19. Testicular tubule. The spermatogenic epithelium has under-
gone a sort of hydropic degeneration. The interstitial con-
nective tissue has become much increased in amount and
has undergone cellular infiltration.
Plate IX.

Fig. 20. Xormal structure of epididymis tubule, x 140.

Fig. 21. Epididymis tubule showing exfoliation of the lining membrane,
and cellular debris in the lumen, x 140.

Fig. 22. Atrophy and degeneration of epididymis tubule. The inter-
stitial connective tissue is much increased in amount, x 140.

Fig. 23. Same, but in more advanced stage, x 140.

Plate X.

Fig. 24. About the same as Fig. 22. x 140.

Fig. 25. ilarked degeneration of epididymis tubule. There is a cellular
infiltration of the interstitial tissue, x 60.
Pr.ATE XL

Fig. 26. Spermatozoon, showing constriction at middle of head. The
head is also somewhat contorted, x 670.

Fig. 27. Spermatozoon. Pear shaped head, x 670.

Fig. 28. Spermatozoon. The head is quite long, and pointed at its
posterior end. x 670.

Fig. 29. Spermatozoon, showing a constriction at middle of the head.
X 670.
Plate XII.

Fig. 30. ^licrocephalic spermatozoon, x 670.

Fig. 31. Spermatozoon. The head is small, and pear shaped, x 670.

Fig. 32. Macrocephalic sperm. The middle piece is nuuli thickened.
X 670.

Fig. 33. Tailless spermatozoa, x 670.

-F/ate /

A/udcU.%

CentrosoTneS
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Discarded. Cutobia'^m.

HecLcL

Liaht Inner bocUj

Gnter/or Vodu-le.
Neck

'Posterior nodule
SbiraL filament
CutobLa^rnic bkeath.

Q-m-iixiar nodlxic.
l7?oolucrLL7n.

PLATE II

Fig. 3.

PLATE III

Fig. 4.

Fig. 5.

PLATE IV

Fig. 6.

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Fig. 7.

PLATE V

Fig. 8.

Fig. 9.

PLATE VI

Fig. 10.

Fig. 11.

PLATE VII

Fijr. li

1-ig. VA.

Fig. 14.

Via. 15.

PLATE VIII

Fig. 16.

Fiff. 17.

Fig. 18.

Fig. 19.

PLATI-: IX

Via. 20.

Fig. 21.

V\ii. 2-2.

Fitr. 2;?.

PLATE X

Fig. 25.

PLATE XI

Fijr. -ilj.

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Fig. 28.

Fig. 29.

PLATE XII

Fig. 30.

Fig. 31.

Fig. 32.

Fig. 33.

i

50777S

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