Book - Uterine and tubal gestation (1903) 1-1

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A personal message from Dr Mark Hill (May 2020)  
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I have decided to take early retirement in September 2020. During the many years online I have received wonderful feedback from many readers, researchers and students interested in human embryology. I especially thank my research collaborators and contributors to the site. The good news is Embryology will remain online and I will continue my association with UNSW Australia. I look forward to updating and including the many exciting new discoveries in Embryology!

Bandler SW. Uterine and tubal gestation. (1903) William Wood & Company, New York.

Uterine and Tubal Gestation (1903): Part I. The Essentials of Uterine Gestation I. The Processes Antedating Uterine Gestation | II. The Embedding of the Ovum in the Guinea-Pig | III. The Embedding of the Human Ovum | IV. The Early Development of the Human Ovum | V. The Trophoblast in the Ova of Animals | VI. The Trophoblast of the Human Ovum | VII. The Further Development of the Human Ovum | VIII. The Chorionic Villi | IX. The Membrana Chorii | X. The Blood-Forming Function of the Trophoblast | XI. The Further Development of the Uterine Placenta | XII. The Placenta | XIII. The Umbilical Vessels and Cord | XIV. Gross Anatomy of the Placenta Part II. The Essentials of Tubal Gestation I. Processes Antedating Gestation in the Tube | II. Varying Views Concerning the Histology of Tubal Gestation | III. Embedding of the Ovum and the Development of Extra-Embryonal structures | Part III. Ovarian and Placental Secretion
Online Editor 
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This early historic 1903 textbook by Samuel Wyllis Bandler (1869-1932) described the understanding of human normal and abnormal implantation at that time. Some of these historic theories described in this textbook have now proved inaccurate or incorrect. Note that all early human developmental stages were still described as the "ovum", today this would be described as the zygote, morula, and blastocyst stages with implantation occurring in week 2.

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Also by this author: The Endocrines (1921)

Modern Notes: implantation | placenta | ectopic pregnancy | Week 2 | blastocyst

Search PubMed: embryo implantation

Historic Disclaimer - information about historic embryology pages 
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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

Part I. The Essentials of Uterine Gestation

Chapter I. The Processes Antedating Uterine Gestation

A. The Trophic Influence of the Ovary

Halban transplanted the ovaries in newly born guinea-pigs to see what effect transplantation and castration would exert upon the development of the genitalia and the breasts. The comparisons with non-castrated animals are very interesting. In the castrated guinea-pig the breasts were later found to be one-fourth the normal size; the genitalia were small, the vulva was one-third smaller than normal ; the uterus was as small as at birth, showing very slight development of muscle and endometrium, and containing no ciliated cells. The breasts showed no glandular tissue, the mammilla? were hypoplastic, the vaginal mucous membrane showed both squamous and cylindrical epithelium — a novel condition. When the ovaries were transplanted under the skin, the uterus was well developed, the tubes were normal, the vulva was large, as were the mammillae likewise. In the transplantation of one ovary a part of the uterus and a piece of the tube had likewise been taken, and these grew in a normal manner, and increased in size. The transplanted piece of uterus contained well-developed muscle and normally secreting glands. The portion of tube showed muscle, epithelium, and ciliated cells. Halban thus showed the influence of the ovary, as a secreting organ, upon the uterus, vagina, vulva, and mamma. He considers it probable that the ovarian secretion acts upon the uterus ; the uterine secretion acting in turn upon the mammary gland.

Ribbert implanted the mamma of a young guinea-pig, with its covering of skin, into a cut near the ear. The wound healed, and five months after the operation, the animal having borne two young, this mamma secreted milk normally, a proof that the connection between the breasts and the ovary and uterus is to be found in no other channels than those of the circulation.

Hegar and Kehrer showed that in newly-born animals, after castration, the uterus does not develop, but remains at the same stage, or even undergoes atrophy, Sokoloff castrated rabbits and dogs and found that, a short time after, they became fat and apathetic, and grew constantly worse. Subsequent examination of the uterus divulged muscular atrophy, especially of the circular layer ; the vessels were fewer, the walls thicker, the lumen was small and even obliterated. The mucosa, however, remained unchanged. When only one ovary had been removed, there ivas no change in the sexual development of these animals, the young ones evidencing a normal and constant development of the genitalia.

Jentzer and Beuttner examined the organs of cows within one year to twenty-two months after castration, and found all the layers of the uterus atrophied and substituted by connective tissue. The stratum vasculare had greatly changed ; the atrophy of the mucosa, while not constant, was in most cases acute. They castrated rabbits, gave subsequently hypodermatic injections of ovarin, and yet the uterus atrophied. The objections to considering this last procedure conclusive are : 1. That the ovarin of cows was used. 2. That it was given hypodermatically. 3. That there was no sufficient increase in the amounts used. The castration of other rabbits showed that the cylindrical epithelium became lower, the protoplasm degenerated, these processes becoming more defined the longer after castration the animals were examined. The muscularis was found to be atrophied ; the larger vessels were gone. Although glands were present, the mucosa showed decided atrophy. In dogs castration produced atrophy of the muscle, but only after three or four months ; the epithelium of the uterus was lower, the protoplasma cloudy, the glands were degenerated, and the vessel walls thickened.

Knauer transplanted the ovaries of rabbits and dogs between the fasciae of the abdominal wall and into the mesometrium, being careful to remove absolutely every bit of ovarian structure. In the mesentery he fastened the ovary with two sutures between two folds of peritoneum, the ovaries being then nourished through endosmosis or through plasmatic circulation. New vessels grew into the ovarian tissue and furnished its subsequent support ; this change began as early as the fourth day. Examination at various periods showed that a small part of each ovary usually degenerated, and new connective tissue appeared in place of the lost cells. In all cases in which a complete degeneration of the ovaries was found, atrophy of the breasts and of the genitalia had occurred. The muscle of the uterus was atrophied, the intermuscular connective tissue was increased, the mucous membrane was atrophied — changes like those which occurred after double castration. Retention of function on the part of the transplanted ovaries was always evidenced by the growth of follicles in a normal manner, by the ripening of the follicles, and by the discharge of the ova. In all such cases the normal character of the breasts, of the uterus, and of the genitalia ivas preserved, and in the younger animals all these organs underwent a natural development. In one dog with well-developed breasts, thirteen months after transplantation an opening of the abdomen showed the right ovary to be of normal size and to contain three follicles. The left ovary was small. Two months later coitus took place, and after two more months two well-developed young were born. Three years after transplantation had been done, this animal on examination showed externally a normal condition of development of the breasts and the genitalia. The right ovary was the size of a pea and contained follicles ; the left ovary was larger than on the previous examination. The right ovary was covered with germinal epithelium; many follicles but no primary follicles being present. A like condition was found in the left ovary. The uterus was normal in every way ; the mucous lining contained many glands and ciliated epithelium. The breasts were of normal character, with normal secretion and glands. Knauer's results proved that the preservation to the organism of the functionating ovaries preserves the breasts, the genital organs, and the sexual instinct, a result possible only through the absorption into the circulation of ovarian secretion, and that the function which the ovary exerts upon the body stands in closest relation to its ability to form ripe ova. Ovarian tissue which has ceased to develop ova has lost its other function, that is, secretion. Therefore the numerous nerves of the ovary are in all probability only vessel nerves. Knauer transplanted in sixteen cases the ovaries of animals into each other, obtaining the above good results in only two cases, since in the others the ovaries degenerated. This shows that a certain relation exists between the cells of one and the same body. Although difficult, transplantation of ovaries from one animal to another is possible.

Morris removed the adnexa in a human patient and sewed a piece of one ovary into the stump of the right tube. One month later the patient became gravid, aborting in the third month. Into a twenty-year-old girl with uterus infantilis and rudimentary adnexa, with suppressio mensium, he transplanted a piece of the ovary of a thirty-year-old patient, fixing it in the fundus of the uterus. Eight weeks later a profuse menstruation, lasting ten days, occurred. ; after six weeks a normal one lasting five days ; the third one five days later. The fourth and fifth menstruations were normal, at intervals of four weeks.

Glass transplanted an ovary of a seventeen-year-old girl into a twenty-nine-year-old patient from whom both ovaries had been removed two years before. The ovary was fixed in its normal situation through an opening in the sac of Douglas, per vaginam. Sixteen days later occurred a bleeding lasting two days ; six months afterward a menstruation of three days ; eight months after operation the patient showed good color, was of good appearance, with a return to' former mental and body conditions, after two years of artificial climax.

Dudley implanted an ovary into the fundus of the uterus after removal of a pyosalpinx duplex, and regular menstruation continued.

An examination of the uterus after castration in women shows an atrophy of the cervix, an immediate atrophy of the corpus, a sclerosis of the vessels which show a growth of the intima, and an endarteritis obliterans, especially in the larger vessels. Few glands are present, and the connective tissue is increased.

Gottschalk found an exceptional condition in one case, in which the muscularis was well preserved, although the mucous lining had almost entirely disappeared.

Cholmogoroff found the uteri at climacterium to contain 9 large amount of connective tissue, the vessels being greatly sclerosed. I have found the same condition in a high degree in two cases of my own.

Normal menstruation is absolute evidence of the presence of a normally functionating ovary. The absence of ovaries, a poor development of these glands, an insufficient secretion of ovarian substance, or a diversion of ovarian secretion to other organs of the body, always causes a failure of uterine development or uterine atrophy. It is possible that, with partial or total absence or atrophy of the uterus or other genital organs, normal ovaries may be present. It is likewise possible for poorly developed ovaries to be the result of a failure of development of the body in general or to be simply a failure of development of the ovary itself, through embryonal disturbances.

Winckel has shown that, in the development of the uterus and tubes, the situation of the Wolffian Body close to the ducts of Muller may influence, to a very great degree, their growth and is a frequent cause of malformation. The early completion of the Wolffian bodies, their opening into the sinus urogenitalis, the growth of the Muller 's ducts along the Wolffian, and their crossing at that spot where the union of Muller 's ducts finds its upper limit, are anatomical embryological factors easily recognized as causes of uterine maldevelopment and hypoplasia. Further, the origin of the ligamentum ileo-genitale rotundum at this upper limit, its close union with the ducts of Muller, the fact that its line of development in a measure opposes the union of the ducts, in addition to tension, pressure, and torsion exerted by the neighboring organs, such as the Wolffian bodies, the bladder, the ureters, the vessels and nerves of the uterus and rectum, are important factors influencing the development of the uterus. In addition, Winckel recognizes the occurrence of abnormal cells in the septum between the ducts of Muller, and general hypoplasia of the vessel system, as additional causes of maldevelopment. Aside from secondary atrophy, the result of constitutional diseases, the embryological cause here mentioned, and the forms associated with general hypoplasia, we recognize in the ovary and its secretion the factor which governs the development of the uterus, the genitalia, and the breasts, and the factor which is concerned in the preservation of these organs and the regulation of menstruation.

Eberlin 1 describes a patient with vaginal defects, who, in her eighteenth year, had vicarious menstruation from the nose at regular intervals for six months, after which no recurrence was noted. Her mammae and external genitalia were small. She suffered from molimina menstrualia every three weeks, at first for periods of three to four days. Eventually severe pain was experienced constantly, associated with sickness at the stomach and vomiting. Laparatomy divulged a uterus rudimentarius, with absence of the adnexa of the left side. The right tube and ovary were almost normal. The uterus was 3.5 centimetres in length, differing microscopically in nowise from the normal as regards muscle, but possessing no cavity. The ovary showed a thickening and a hyaline degeneration of the vessel walls. In addition to the corpus luteum, very few follicles but no Graafian follicles were present. After castration all the annoying symptoms disappeared.

Fritsch operated in a like case in which severe molimina men^ineberg, strualia, sickness at the stomach, and rectal bleedings were the indications. Am. Jour, of Obstet., No. 4, 1898.

In all, Eberlin finds twenty-one cases of this sort associated with vagina] defect to have been operated upon. The ovary in the above-cited case of Eberlin, though containing few follicles, nevertheless secreted sufficiently to cause the severe symptoms. Since no endometrium was present, and no excretion by that channel took place, the ovarian secretion, through cumulative action, was responsible for the severity of the pain. Whatever part the uterus took in these attacks must have deen the result of uterine contraction, since congestion alone is not responsible for so much pain, as is evidenced by the absence of such suffering in pure congestions of the uterus, associated with metritis, etc.

B. Constitutional Changes Dependent on the Ovary

Were we to consider the relation between the ovary and the uterus, and between the genitalia and other organs to exist through the medium of the cerebro-spinal or sympathetic systems, why should we find the walls of the tubes thin and containing more connective tissue in cases of poorly developed ovaries and at the menopause, when no primary follicles and no Graafian follicles are present in the ovaries? The uterus then shrinks, the walls become thin and flabby, the mucous membrane atrophies, the connective-tissue elements are more prominent, and the ciliated epithelium disappears. Is it not extraordinary to make the occurrence of these decided changes dependent upon a cessation of ovarian congestion? That these differences are not due to the lack simply of reflex stimulation on the part of the ovary is sufficiently evidenced by the constitutional changes occurring at puberty, at menstruation, during pregnancy, and at the menopause. The relation between the changes occurring at puberty and at the menopause, before menstruation and after menstruation, during pregnancy and after pregnancy, show a decided resemblance. Until shortly before each menstrual period, temperature, pulse, muscular activity, lung capacity, and the excretion of urea increase, and reach their maximum two to three days before the appearance of blood. During this period we find hyperemia, edema, increased activity of the ovary, changes in all the mucous membranes, and increased function of the glandular apparatus. The occurrence of swelling of the breasts, tenderness of the abdomen, even pain and the passage from the vagina of increased mucus, sometimes mixed with blood, prove at the beginning of each menstrual period a wave movement and an increased blood tension by no possibility due to reflex causes. During and after menstruation regressive changes are evident, and the excretion of nitrogenous elements is diminished.

During pregnancy we have an increased amount of the watery elements of the blood, an increased proportion of fibrin, a diminished amount of albumin, an increase in the white blood cells, a genuine increase in the number of red blood cells and in the amount of hemoglobin.

Before labor the temperature is highest in the last three months of pregnancy, and there is an increase in the elements of the body, equal to one-thirteenth of the body weight. This increase is due to serous infiltration, and to the increased ability of the body to form organized tissue. Post partum, after a temporarily short rise, the temperature is lower, the blood pressure sinks, and becomes normal on the ninth day. The loss in body weight is equal to one-ninth of the weight of the pregnant person. For labor and post partum together, there is a loss of weight nearly equal to one-fifth of the body weight at full term. After labor there is a diminution of tissue change and a diminution in the amount of urine.

Therefore as regards temperature, blood pressure, body weight, the amount of urine secreted, etc., there is always a similar increase before menstruation, and a like decrease in intensity during and after menstruation, as during and after parturition, so that Virchow has well characterized menstruation as being a labor en miniature.

According to Loewenthal, who believes that every ovum is embedded in the mucous membrane of the uterus, we may say that labor is a menstruation in which a fully developed ovum is expelled. The coincidence of labor with a menstrual period seems therefore natural. The resemblance of the mechanical processes concerned in menstruation and in labor is remarkable, and these changes can be due only to the stimulative effect of the ovarian secretion, acting not alone upon the uterus and the genital organs, but likewise upon the breasts, upon the blood elements, and upon blood tension. This secretion is a stimulus likewise to uterine contractions, and is the probable cause of the contractions normally occurring during pregnancy. The action of this accumulated secretion upon the uterus at the end of pregnancy is probably the cause of labor pains.

The action of the ovarian secretion upon pulse tension, and its effect upon the mucous membranes of the body generally, are likewise evidenced by the congestion of the vocal cords during menstruation, so that during this time the singing voice is poor. The secretion of intestinal mucus is also greater, there is increased perspiration, the lower turbinated bones are swollen, and the eye suffers limitations in power. The best evidence of the constitutional elements involved in the process of menstruation is the occurrence of vicarious menstruation. Under this designation ice consider bleedings occurring at regular intervals in a patient suffering from amenorrhea. The most frequent spot for this bleeding is the nose, usually the lower turbinated bones.

Fleiss described a two-year-old child with well-developed pubes and breasts who had periodical bleedings from the nose. In a second case, a girl fourteen years old, well developed, suffered from regular bleedings from the nose, which stopped, however, Avhen real menstruation began. In another case the regular nose bleedings at intervals of twenty-nine days stopped during pregnancy, only to begin afterward, and after a continuation of eight months ceased again on the occurrence of a second pregnancy. Bleedings of the same kind have been described as occurring regularly from other mucous membranes, the trachea, the lungs, and the stomach. In the latter instance the bleedings were not always associated with vomiting, the blood being usually found in the feces. In other cases there were bleedings into the thyroid gland. In cases with poorly-developed uteri, these bleedings disappeared only when the uterus began to functionate properly.

The secretion of the ovary exerts through the medium of the blood a stimulating effect upon the breasts, noticeable before menstruation, during pregnancy, and during the period of lactation .

(ioltz cut through the cord of a dog at the level of the first lumbar vertebra, and later saw the signs of rut appear. After coitus one dead and two living young were born. The breasts were well developed, and lactation and nursing followed the normal course. Since these changes, the sexual tendency and the process of labor could not have been excited through the cord, it must be that a certain secretion of the ovary, acting through the medium of the circulation, gives the stimulus for the exercise of those functions.

Ovarian secretion exerts a most decided effect upon the development of the uterus and the genitalia, as well as upon the breasts. It is absolutely necessary for the preservation of the developed uterus, the other genitalia, and the breasts. An overproduction of ovarian secretion or a disturbance in the function of the ovary causes pathological changes ; an under-production of ovarian secretion is likewise the cause of pathological conditions. The production of ovarian secretion and its action for a certain period of time are the probable causes of the pains of labor.

C. Menstruation

Menstruation is the periodical loss of blood from the uterus or from any mucous membrane, occurring for the reason that no fecundated ovum is present in uterus or tube, and may be divided into three periods : the premenstrual, the menstrual, and the postmenstrual. During the premenstrual period, the ten days immediately preceding the appearance of blood, the following changes take place :

The superficial capillaries become greatly dilated, and serous infiltration of the endometrium takes place, which separates the meshes of the stroma, accompanied by a gradual but decided dilatation of all the blood vessels and lymph channels. There occur a growth of round cells in the interglandular tissue, and an entrance of leucocytes into the mucous membrane. The glands become larger and wider, being often filled with secretion.

This swelling of the mucous membrane, the dilatation of the blood vessels, the production of round cells, and the growth of the superficial layer of the endometrium produce the so-called decidua menstrualis. Although in the connective tissue large cells, not to be distinguished from young stages of decidua cells, are found, it is to be noted that typical decidua cells do not, as a rule, develop at this time in the superficial layer. The endometrium is at this period from 6 to 7 millimetres in thickness.

The period during which blood is thrown out is the next, or menstrual period. The superficial capillaries are greatly dilated, and an exit of blood elements, not dependent on a bursting of the capillaries, goes on for several days. The bleeding occurs partly through diapedesis, and, in strong bleedings, through rhexis. There is little or no destruction of the mucosa, only a very slight fatty degeneration of the epithelium of the uppermost layer, so that in the excreted blood relatively few epithelial cells are found.

After menstruation, the uterus shows an almost continuous covering of epithelium, interrupted in spots, especially if pathological processes be present. Most of the ciliated epithelium is preserved. The first stimulus to bleeding is due to contraction of the uterus, which at the height of congestion is possibly accompanied by contractions of the tube. During menstruation the uterus is larger and in the first few days following likewise soft and flabby. The flabbiness lasts longer than the bleeding. A spontaneous dilatation of the cervix canal takes place, and reaches its height on the third or fourth day. This dilatation takes place without regard to the amount of blood discharged, whether the menstruation be painful or painless. The cervix is hyperemic, the glands showing an increased secretion of mucus. The blood thrown off is mixed with the mucus of the uterus and cervix, and later with the acid secretion of the vagina, for this reason coagulating less easily than other blood.

The secretion from the sebaceous glands of the external genitalia is responsible for the peculiar odor present in some cases, and in nowise differs from that observed in the axilla. On the inverted uterus the blood has been observed to appear in single drops from the region of the openings of the glands. The endometrium is then covered with white and bloody mucus, the openings of the glands being clearly seen. There is no degeneration or destruction of the mucous membrane. The changes in the tubes, if any, are slight (Gusserow).

The next, or postmenstrual period comprises fourteen days, during which the mucous membrane returns to a thickness of 3 millimetres. During this time all cells not capable of further growth are thrown off, and the epithelium, only partially denuded, is regenerated. The epithelial cells found singly or in groups in the lumen of the glands disappear by resorption, and partially by phagocytosis. From the sixth day after the beginning of menstruation, cell division is prominent. It is probable that mitosis takes place, even during the exudation of blood. The presence of mitosis in the leucocytes is an evidence of growth of the interstitial tissue during menstruation.

Regeneration concerns the stroma, the glands, and surface epithelium, reaching its height on the fourteenth or fifteenth day after the beginning of menstruation. The mucous membrane becomes pale, the glands and vessels have returned to their normal state, the lost epithelium has been restored. From the middle of the third week on there is a diminution in mitosis.

Menstruation is not a process by which the mucous lining of the uterus is thrown off, with subsequent regeneration previous to the next menstruation. It is simply the excretion of blood from the decidua menstrualis occurring for the simple and sole reason that there is in the uterus no fecundated ovum.

Ovarian secretion is the direct cause of this periodical swelling of the mucosa, and it continues, if fecundation has taken place, to exert, not alone a local, but likewise a stimulating influence on the general and sexual organs. This stimulation occurs to a heightened degree during pregnancy, and on removal of the uterus, since the secretion is ordinarily excreted during menstruation. On the occurrence of pregnancy, this secretion causes a further development of the uterus and the decidua, and plays an important part in the process of labor; it stimulates the functions of the breasts, exerts a decided constitutional stimulation, and is the cause of many of the pathological conditions occurring before and during labor.

The generally accepted view as to the relation between ovulation and menstruation is that of Pfliiger. The increase of the contents of the Graafian follicle, which results through the secreting activity of the cells of the membrana granulosa, is said to stimulate the nerves running in the stroma and ending in the cells. The increasing tension reflexly irritates the vasomotor centre, with resulting dilatation and congestion of the uterine vessels. At the height of this congestion the follicle is supposed to burst, through pressure. As congestion and tension in the ovary are supposedly the reflex causes of the congestion of the uterus and pelvic organs, menstrual bleeding is considered a result of periodical ovulation. This theory obtained support through the experiments of Strassman, who showed that increased tension in the ovaries of animals, produced by the injection into them of fluid, caused swelling of the mucous membrane of the uterus. In answer it may be asked why the same condition does not result in all cases of ovarian tumors and ovarian cysts, which may develop entirely beneath the albuginea.

This theory of Pfliiger and the experiments of Strassman have been generally quoted to prove the causal relation of ovulation to menstruation. The congestion in the ovary, and the swelling of the Graafian follicle, must then, through the nervous system, stimulate the uterus to congestion, with a resulting periodical bleeding. The cessation of menstruation after castration has likewise been cited as proof of the fact that without ovulation there could be no menstruation, since the periodical hyperemia reflexly caused by ovulation no longer takes place. By others it was considered that the distension of the ovary, occurring periodically, exerted an action upon the vasomotor system, then causing congestion. The atrophy of the uterus after castration was likewise pointed out as a proof of the resulting diminished Mood supply, ordinarily furnished by this reflex action of the ovary. On the other hand, it has been frequently stated that the result of castration was due to the tying off of the arterise spermatica? internas, and that the resulting anemia was the cause of atrophy of the uterus, and of the diminution in the size of the uterine myomata.

Rein cut all the sympathetic and all the spinal nerves running to the uterus in a dog, and yet the animal bore young. Inasmuch as the uterus, freed from the central nervous system, performed its functions as before, he concluded that a regulating nerve centre exists in the ovaries. All this was taken to prove that menstruation was a result of reflex action originating in the ovaries.

D. The Action of Ovarian Secretion on the Endometrium

The periodical swelling of the mucous membrane is due to the secretion given of by the ovary, and the experiment of Strassman proves that the forcing of an additional amount of this secretion into the circulation only enhances this effect. That ovulation and menstruation, or at least the latter, occur with a certain regularity is a fact which we are not able to explain any more than we can say why in malaria the plasmodia are thrown into the circulation at regular intervals, or why the menopause usually occurs at a certain period. At any rate, were menstruation the direct result of periodical ovulation, we should still be in need of a satisfactory explanation for the latter phenomenon. No other theory is satisfactory than that of secretion. Ovulation and menstruation are evidences of the functional capability of the ovary. Since ovulation may occur without menstruation, but the latter never without the former, we have here an evidence that a certain functional activity of the ovary is necessary to stimulate the mucous membrane to its periodical changes. That after double castration regular bleedings may occur for a certain period is proof of the independent role which the endometrium, to a certain degree, plays in the process of menstruation, for sufficient of the secretion of the ovaries may still remain in the circulation to produce the normal processes after removal of the ovaries. The part which the ovaries play in the development of the body, the effect of their presence upon the breasts and the genital tract at puberty, before each menstrual period, at the menopause, and after castration, are sufficient evidence of their secreting power. The experiments of Knauer and others prove that it is simply the presence of the ovaries, and the preservation of their secretion, which are of importance to the body, and that their action upon the uterus is in nowise reflex in character, since when removed and implanted elsewhere, and in this way loosened from their connection with nerve plexuses and the nervous system, every sexual peculiarity is absolutely preserved.

Menstruation is simply an evidence that a fecundated ovum is not present in the tube or in the uterus. Practically no part of the endometrium is thrown off. It seems as if the blood thus lost simply gave an exit to the secretion of the ovary, which, if retained too long in the body, produces in pregnancy certain normal changes, and in a large number of cases abnormal processes. In pregnancy the ovarian secretion stimulates the uterus to enlargement and growth, it stimulates the formation of blood, produces tension and congestion in the vascular system, and stimulates the function of the breasts. After labor, lactation is stimulated by this secretion, so that little or no effect is exerted upon the uterus ; therefore it rarely undergoes the periodical stimulation and has a tendency to atrophy. The changes occurring in the decidua menstrualis and the decidua graviditatis are certainly evidences of the action of a secreted substance. The method in which the ovum is embedded in the decidua, and the processes occurring in its immediate vicinity, as well as the fact that in extrauterine gravidity a decidua graviditatis is formed in the uterus, with decided enlargement of this organ, are likewise proofs of this fact.

E. The Relation of Ovulation and Menstruation

As a matter of fact, ovulation and menstruation are related only in that both are the result of the secreting function of the ovary, and are in nowise connected as regards cause and effect. The ovum is the external secretion, the internal secretion entering the blood through the lymph channels. The ripening and expulsion of an ovum may occur at any time. It is, however, a fact that menstruation occurs only after the ovary is capable of producing ripe ova, and whether the egg be expelled, or occasionally degenerate in the follicle, is immaterial. The independence of ovulation and menstruation is evidenced by the fact that the former takes place before menstruation has appeared, and likewise after the menopause, as is proven by cases, by no means rare, of pregnancy in girls who have not yet menstruated, as well as by the occurrence of pregnancy at variable periods after the amenorrhea of the climacterium. In children the primary follicles develop fully before puberty, and the same development occurs even in the fetus and in the newly born. Those ova and follicles go through the same stages of development an. I ripening as in the case of adults. That they are not capable of fecundation is shown by the fact that the ova are only one-half as large as in adults. In those cases, however, of young children with well-developed breasts and genitalia, when menstruation begins there is an unusually strong development of the body, and the ova, as well as the follicles, differ in no way from those found in menstruating adults.

Leopold found that in forty-two pairs of ovaries, thirty pairs showed a corpus luteum belonging to the last menstruation. In thirteen pairs no follicles were found which, on account of their size or the swelling of the follicle, justified a belief that a bursting of any follicle would have occurred at the next menstrual period. In twelve cases no corpus luteum was found belonging to the last menstruation. One case showed a follicle which had burst between menstrual epochs.

Arnold found that in fifty-four cases, only thirty-nine showed the presence of fresh corpora lutea after the last menstrual period.

Williams found this to be the case in twelve cases out of sixteen. Therefore, ovulation, although in the majority of cases occurring at or near the menstrual period, is not the cause of the same.

Leopold has shown experimentally that in the inter-menstrual period follicles ready to burst are present, and that through certain causes, such as coitus, the exit of an ovulum may result at any time. That ovulation occurs during pregnancy is proved in the case mentioned by Kroenig, in which conception occurred as a result of coitus four days post partum. The relatively frequent occurrence of pregnancy during the temporary amenorrhea of lactation is a proof of ovulation during this time.

Consentino finds that during pregnancy and lactation ovulation continues. Although the ovum of a menstrual period may be fecundated immediately thereafter, the ovum usually fecundated is the one given off between four and eight days before menstruation, the egg thus belonging to the period, so to say, first omitted. The ovum retains its vitality for an average period of twelve days, and yet out of a collection of two hundred and fourteen pregnant cases, in sixty- five coitus had taken place after the twelfth day following menstruation. Therefore, in these pregnancies at least, the ovum must have belonged to what may be called the next awaited menstrual period. Even if the expulsion of the ovum occur only a few days before that time, its fecundation is not difficult to explain, nor are the cases puzzling in which one coitus directly after menstruation produces pregnancy, as the spermatozoa retain their vitality for a long time, and have been preserved, in proper temperature, for a period of eight days. Duhrssen has found spermatozoa in the tube three and one-half iveeks after the last coitus.

It is therefore evident that ovulation, as a rule, occurs from four to eight days before menstruation, but it may occur at other periods, as ripe ova, practically speaking, may be present at almost any time.

F. Ovulation

The other of the functions of the normal ovary is the production and expulsion of ova capable of being fecundated. It is probable that after birth no new ova are formed from the germinal epithelium. At and after puberty we judge the vitality of the ovary by its ability to bring these ova to a stage which may be called ripe. For the expulsion of an ovum from the Graafian follicle, a gradual increase in size of the follicles takes place, depending partly on an increase in the amount of liquor folliculi. The cells of the follicle epithelium undergo fatty degeneration, and the internal layer, the tunica interna, shows an increase in the size of the cells, and a decided development of the blood vessels. The protoplasm enlarges greatly, begins to take on a yellow color, and these cells, now lutein cells, are arranged in several layers, forming an irregular surface. As a result of the fatty degeneration of the follicle epithelium, the ovum is freed from the cumulus oophorus. The most prominent point of the follicle is poor in blood supply, and furnishes the so-called stigma folliculi. It is here that the opening takes place which serves as an outlet for the ovum. This opening is probably the result of the reaction or chemical effect produced by the ripe ovum, since in the newly born, and in children, follicles of the same size and even larger ones exist without bursting — the so-called atresic follicles. The facts that large follicle cysts occur in the ovary without opening, and also that in women in whom the unopened follicles degenerate, disturbances of menstruation occur — so-called missed ovulation — speak for a chemical reaction as probably one of the functions of the ripe ovum. That the orgasmus venereus does play a part in hastening the expulsion of the ovum is not to be questioned. In most cases the opened follicle is then filled with blood which likewise empties into the peritoneal cavity, and the so-called corpus luteum spurium results. If pregnancy takes place, this body develops decidedly, forming the corpus luteum verum (Nagel).

After ovulation, the ovum is thrown out into the abdominal cavity, and then, influenced by the wave movement of the ciliated epithelium of the tubes, the fimbria? of the ampulla, and the fimbria? ovarica?, finds its way into the uterus. It is not necessary that the tube should grasp or surround the ovary, since it would embrace, even under favorable circumstances, only part of the ovary. This wave movement of the ciliated epithelium causes a current in the peritoneal plasma, which directs the ovum into one or the other of the tubes.

Lode injected, with a needle pointed toward the diaphragm, the eggs of ascarides into the abdominal cavity of rabbits, in the region of the umbilicus. After ten hours, without the excitement of coitus, he found these eggs in the tube. An interesting proof of the existence of this current is given by Knauer, who removed the ovaries of dogs, and sewed them into the mesentery. In one case, on opening the abdomen he saw in one of the ovaries three ripe Graafian follicles. Shortly after, coitus took place, and in due time two young were born. A further proof is found in the experiments of Leopold, who showed that an ovum given off by one ovary may enter the tube of the other side. The cases are not rare in which the tube of one side was closed or absent, and although the corpus luteum verum was found in the ovary of the same side, yet the ovum was found in the uterus. The same is true of those cases in which the corpus luteum verum is on one side, and the ovum develops in the other tube, or in the horn of a uterus unicornis of the opposite side. Therefore it is certain that, normally, a wave current exists on either side of the uterus, and that the ovum is attracted by the stronger current, usually that of the side from which it came, possibly, however, by the current of the other side.

It is therefore evident that an ovum, by natural means, may at any time enter the tubes, and that no congestion of the tube, no erection of the tube, and no reflex spasmodic grasping of the ovary by the tube are necessary.

Lode has shown that an ovum finds its way through the tube into the uterus in thirty hours.

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Uterine and Tubal Gestation (1903): Part I. The Essentials of Uterine Gestation I. The Processes Antedating Uterine Gestation | II. The Embedding of the Ovum in the Guinea-Pig | III. The Embedding of the Human Ovum | IV. The Early Development of the Human Ovum | V. The Trophoblast in the Ova of Animals | VI. The Trophoblast of the Human Ovum | VII. The Further Development of the Human Ovum | VIII. The Chorionic Villi | IX. The Membrana Chorii | X. The Blood-Forming Function of the Trophoblast | XI. The Further Development of the Uterine Placenta | XII. The Placenta | XIII. The Umbilical Vessels and Cord | XIV. Gross Anatomy of the Placenta Part II. The Essentials of Tubal Gestation I. Processes Antedating Gestation in the Tube | II. Varying Views Concerning the Histology of Tubal Gestation | III. Embedding of the Ovum and the Development of Extra-Embryonal structures | Part III. Ovarian and Placental Secretion

Reference: Bandler SW. Uterine and tubal gestation. (1903) William Wood & Company, New York.

Cite this page: Hill, M.A. (2020, July 8) Embryology Book - Uterine and tubal gestation (1903) 1-1. Retrieved from

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