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derived from the s % troma of the ovary, and is separable into two parts, an inner
containing the ramifications of the capillary blood-vessels, which are abundantly
distributed to the larger follicles, but nowhere penetrate amongst the epithelium
cells ; and an outer part more fibrous, in which the larger branches of the blood-
vessels of the follicle run. In both layers of the follicular wall, the cells are similar
to those of the general stroma, interstitial cells occurring abundantly ; but it is
uncertain whether there are any cells present of the nature of muscular fibres. The
smaller blood-vessels running round the follicle from its deepest part, and minutely
sub-divided on its inner surface, converge towards a point near the middle of the
most projecting part, called the stigma. This marks the spot where the rupture
of the vesicle ultimately occurs, when fully matured.

Each Graafian follicle usually contains only a single ovum ; but occasionally,
though seldom, two ova, and very rarely three, are observed in the same follicle.
The structure of the ovum has already been described in Vol. I., pt. 1 ; but for
the sake of completeness the description of the human ovum may be here repeated.

Structure of the ovarian ovum. The human ovum resembles that of all other
mammals (with the exception of monotremes) in its minute size. Immediately
before the time of its discharge from the Graafian follicle of the ovary in which it
has been formed, it is a small spherical vesicle measuring about T |yth inch ('2 mm.)
in diameter, and is just visible as a clear speck to the naked eye. "When
examined with the microscope, it is found to be invested by a comparatively thick,
clear covering. This, when the centre of the ovum is exactly focussed, has the
appearance in optical section of a clear girdle or zone encircling the ovum (fig. 288),
and was hence named zona pellucida by von Baer (1827). But on more careful
examination with higher magnifying powers, and especially by the examination of
sections, there is not much difficulty in making out the existence of striae passing
radially through the membrane (fig. 289, zp). On this account, and especially since a
similar radially striated membrane forms a characteristic part of the investment of
the ovum in many animals belonging to widely different classes, it is more convenient,
in place of the name zona pellucida, which has been exclusively used to designate



this investment in mammals, to employ the more general term zona radiata, or to
speak of it simply as the striated membrane of the ovum.

The zona radiata of the mammalian ovum is sufficiently tough to prevent the
escape of the contents of the ovum, even when subjected to a considerable amount
of pressure. If, however, the pressure be excessive, the tunic splits, and the soft
contents are extruded (fig. 288, I). The striee in the membrane are believed to
be minute pores, and are supposed, while the ovum is yet within the Graafian
follicle, to permit the passage of granules of nutrient material into the interior of
the ovum. After the ovum is discharged from the follicle, the spermatozoa may
perhaps find their way into the ovum through these pores. According to Retzius

Fig. 288. OVARIAN OVUM OF A MAMMAL. (Allen Thomson.)

a, an entire ovum, viewed under pressure ; the granular cells have been removed from the outer
surface, the germinal vesicle is seen in the yolk substance within ; b, the external coat or zona burst by
increased pressure, the yolk protoplasm and the germinal vesicle having escaped from within ; c, germi-
nal ves ; .cle more freed from the yolk substance. In all of them the macula is seen.


sp, zona pellucida, showing radiated structure ; vi, vitellus, round which a delicate membrane is
seen ; <jv, germinal vesicle; ys, germinal spot.

the protoplasm of the ovum is united with the follicle-cells by fibres which pass
through the pores of the zona.

Immediately surrounding the zona radiata, as the ovum lies within the mature Graafian
follicle, is a thin stratum of granular substance, probably deposited upon the exterior of the
ovum by the innermost cells of the discus proligerus, which immediately encircle the ovum
within the follicle. When the Graafian follicle bursts and the ovum is set free, this granular
material appears to imbibe water, and, as is specially noticeable in the ovum of the rabbit,
swells up into a clear gelatinous envelope, which has been termed, from a possible homology
with the white of the bird's egg, the albumen. But in the mammal this structure has not
the nutritive importance to the embryo which is possessed by the corresponding formation
in the bird, and it disappears during the passage of the ovum down the Fallopian tube.

The substance of the ovum within the tunica radiata is known as the vitellus
or yollc (fig. 289, vi). It is a soft semi-fluid substance, composed mainly of proto-
plasm, which is filled with globules and granules (yolk-granules) of different
sizes, but all small, and possessing a high index of refraction. Examined in the
fresh condition, the protoplasm between the granules looks perfectly clear and
structureless, but after treatment with suitable reagents, it may be seen to consist of
a fine reticulum, which is especially fine and close near the periphery of the ovum,
and also around the germinal vesicle, at which places the yolk-granules are in less
amount than elsewhere. The substances which occur within an ovum other than



the nucleus and protoplasm, may, as in cells generally, be collectively designated
*' deutoplasm " ; they are regarded as furnishing a supply of nutrient matter to the
protoplasm during the earlier stages of development.

Embedded in the protoplasmic vitellus, usually eccentrically, is a large spherical!
nucleus, which was termed by its discoverer, Purkinje, the germinal vesicle? This,,
which is about -g^th inch ('05 mm.) in diameter, has all the characters of a nucleus
of a cell. It consists of a nuclear membrane (fig. 289, gv} enclosing a clear material
or matrix, embedded within which may be seen strands of karyoplasm, enclosing-
one or more well-marked nucleoli. Frequently there is but one nucleolus, which is
then large and prominent, and has received the name of germinal sjtot (macula ger-
minativa, Wagner, 1835).

There is some doubt whether, before fertilization, there is another membrane (vitelline
membrane) enclosing the vitellus within the zona radiata. The evidence of the presence of
such a membrane is by no means clear, although its existence has been maintained by very
competent observers (v. Beneden, Balfour).

Structure of the corpora lutea. The corpora lutea are produced after the
rupture of the Graafian follicles and the escape of their contents by what may


A, commencing ingrowth of the vascular tissue of the theca f olliculi into the hvpertrophied follicular
epithelium ; , vascular ingrowth ; thi, theca or wall of follicle ; fe, follicular epithelium.

B, a further stage in which the vascular ingrowths of the theca converge towards a central cavity.
Between the ingrowths or trabecul* the follicular epithelial cells, which are undergoing rapid
multiplication, appsar as if disposed in columns. I, leucocytes amongst the follicular cells ; ke, surface
epithelium of the ovary.

c, a further stage, the columns being now narrower and the trabeculse more numerous.

1 Purkinje discovered the germinal vesicle in the bird's ovum in 1825 ; that of mammals was first
noticed by Coste in 1833.



perhaps be most correctly described as a process of hypertrophy, i.e., growth of the
walls of the empty follicles. The hypertrophied follicular wall becomes thrown into
plaits or folds which as they increase in extent occupy more and more of~the cavity
of the empty follicle, until this has become entirely filled. The hypertrophy is
usually described as the result of the proliferation of the polyhedral interstitial
stroma-cells, which as already stated occur in the wall of the follicle in abundance,
and there is in addition a considerable development of blood-vessels, which run,
accompanied by fibrous tissue, into the folds into which the wall of the follicle is

PLETED. (Sobotta. )

The central cavity is now occu-
pied by jelly-like connective tissus
and the converging trabeculffi anas-
tomose with one another so as some-
what to break up the columnar
arrangement of the luteal cells.

thrown, giving off capil-
laries which ramify abun-
dantly in the folded wall.
But according to the obser-
vations of J. Sobotta upon
the mouse, the main part
of the thickening is due to
a simple hypertrophy of
the epithelium-cells of the
membrana granulosa, into
which vascular processes of
the wall of the follicle grow

and amongst which a certain number of leucocytes penetrate (fig. 290). Meanwhile
the irregular cleft-like space which now alone represents the cavity of the follicle, as
well as the opening resulting from the rupture of the follicle and by which its cavity
communicated with the surface, become occupied by a sort of jelly-like connective
tissue, which constitutes a kind of hilum for the follicle. To this central fibrous
band the strands of fibrous tissue which accompany the blood-vessels in the folds of
the hypertrophied wall of the corpus luteum converge. At the same time the plaited
disposition of the wall becomes in great measure obscured, so that a section of a
corpus luteum, when advanced in development (fig. 291), exhibits a fibrous frame-
work having a radial disposition, with the intervals between the radiating trabeculge
occupied by a tissue which is almost wholly composed of large yellowish cells.
Amongst these cells are numerous cleft-like spaces (lymphatic), and except for the
fact that the columnar disposition is less distinct and that the capillary blood-
vessels come more closely into relationship with the cells of the tissue, the structural
appearances are not unlike those which are met with in the cortical part of the
suprarenal capsule.

The corpus luteum is at first sharply marked off by the theca folliculi
from the surrounding ovarian stroma, but after a time its limits are less well
defined from the neighbouring parts of the stroma, into which it may be said
gradually to merge and in this way to disappear. The result is that, as age
advances, the stroma of the ovaries, at least in some animals, becomes gradually
pervaded with cells like those of the corpora lutea.

Vessels and nerves of the ovaries. Arteries. The ovaries are most directly



supplied by the ovarian arteries, analogous to the spermatic in the male, which
anastomose freely by an internal branch with the termination of the uterine arteries.
Sometimes this anastomotic branch is so large that the ovary seems to be supplied
almost entirely by the uterine artery. The ovarian artery always sends numerous
branches to the Fallopian tube. The smaller arteries penetrate the ovary along its
attached border, pierce the proper coat, and run in flexuous parallel lines through its
substance. The veins correspond, forming a plexus near the ovary named thepam-
piniform plexus. The nerves are derived from the ovarian plexus ; and also from the
uterine nerves, which invariably send offsets to the Fallopian tubes. The nerves are


SEXES (from Farre, after Kobelt).


a, a, Epoophoron (parovarium) formed from the upper part of the Wolffian body ; b, remains of the
uppermost tubes, sometimes forming hydatids ; c, middle set of tabes ; d, some lower atrophied tubes ;
e, atrophied remains of the Wolffian duct ; /, the terminal bulb or hydatid ; A, the Fallopian tube,
originally the duct of Miiller ; i, hydatid attached to the extremity ; I, the ovary.


a, a, convoluted tubes in the head of the epididymis developed from the upper part of the Wolffian
body ; b and f, hydatids in the head of the epididymis ; c, coni vasculosi ; d, vasa aberrantia ; h,
remains of the duct of Muller with i, the hydatid of Morgagni, at its upper end ; Z, body of the testis.

said by Gawronsky to penetrate even amongst the epithelium cells of the membrana
granulosa of the Graafian follicles.

Parovarium. The organ so named by Kobelt, or the organ of Roscnmuller,
who first described it, is a structure which can usually be brought plainly into
view by holding against the light the fold of peritoneum between the ovary
and Fallopian tube (see fig. 292, a, b, c, d). It consists of a group of
scattered tubules lying transversely between the Fallopian tube and ovary, lined
with epithelium, but having no external openings. The tubules converge towards
their ovarian end, but remain separate there, while at the other they are more or
less distinctly united by a longitudinal tube which is sometimes of considerable size,
and prolonged for some distance downwards in the broad ligament. Its more
developed form in some animals, as the cow and pig, constitutes the duct of
Gartner. The origin of this vestige of a foetal structure is referred to under Develop-
ment (Vol. I. part 1, p. 120). Here it is sufficient to state that it corresponds
essentially to the epididymis of the male. Vestiges corresponding to the organ of
Giraldes of the male are also sometimes to be detected in the adult female, in the
shape of tubular remnants, situated in the broad ligament nearer to the uterus
than the parovarium. These constitute the paroophoron of Waldeyer ; the organ of
Rosenmiiiler constituting the epoophoron.



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The mammary glands (mammce), which yield the milk in the female, are accessory
parts to the reproductive system. They give a name to a large class of animals
(Mammalia) which are distinguished by the possession of these organs. When
fully developed in the human female, they form, together with the integuments and
a considerable quantity of fat, two hemispherical or conical eminences (the breasts)

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