The Evolution of Man — Volume 2

Chapter 17

Chapter 175,355 wordsPublic domain

In the Selachii also we find a longitudinal row of segmental canals on each side, which open outwards into the primitive renal ducts (nephrotomes, Chapter 1.14). The segmental canals (a pair in each segment of the middle part of the body) open internally by a ciliated funnel into the body-cavity. From the posterior group of these organs a compact primitive kidney is formed, the anterior group taking part in the construction of the sexual organs.

In the same simple form that remains throughout life in the Myxinoides and partly in the Selachii we find the primitive kidney first developing in the embryo of man and the higher Craniotes (Figures 2.386 and 2.387). Of the two parts that compose the comb-shaped primitive kidney the longitudinal channel, or nephroduct, is always the first to appear; afterwards the transverse "canals," the excreting nephridia, are formed in the mesoderm; and after this again the Malpighian capsules with their arterial coils are associated with these as coelous outgrowths. The primitive renal duct, which appears first, is found in all craniote embryos at the early stage in which the differentiation of the medullary tube takes place in the ectoderm, the severance of the chorda from the visceral layer in the entoderm, and the first trace of the coelom-pouches arises between the limiting layers (Figure 2.385). The nephroduct (ung) is seen on each side, directly under the horny plate, in the shape of a long, thin, thread-like string of cells. It presently hollows out and becomes a canal, running straight from front to back, and clearly showing in the transverse section of the embryo its original position in the space between horny plate (h), primitive segments (uw), and lateral plates (hpl). As the originally very short urinary canals lengthen and multiply, each of the two primitive kidneys assumes the form of a half-feathered leaf (Figure 2.387). The lines of the leaf are represented by the urinary canals (u), and the rib by the outlying nephroduct (w). At the inner edge of the primitive kidneys the rudiment of the ventral sexual gland (g) can now be seen as a body of some size. The hindermost end of the nephroduct opens right behind into the last section of the rectum, thus making a cloaca of it. However, this opening of the nephroducts into the intestine must be regarded as a secondary formation. Originally they open, as the Cyclostomes clearly show, quite independently of the gut, in the external skin of the abdomen.

(FIGURE 2.395. Primitive kidneys and germinal glands of a human embryo, three inches in length (beginning of the sixth week), magnified fifteen times. k germinal gland, u primitive kidney, z diaphragmatic ligament of same, w Wolffian duct (opened on the right), g directing ligament (gubernaculum), a allantoic duct. (From Kollmann.))

In the Myxinoides the primitive kidneys retain this simple comb-shaped structure, and a part of it is preserved in the Selachii; but in all the other Craniotes it is only found for a short time in the embryo, as an ontogenetic reproduction of the earlier phylogenetic structure. In these the primitive kidney soon assumes the form (by the rapid growth, lengthening, increase, and serpentining of the urinary canals) of a large compact gland, of a long, oval or spindle-shaped character, which passes through the greater part of the embryonic body-cavity (Figures 1.183 m, 1.184 m, 2.388 n). It lies near the middle line, directly under the primitive vertebral column, and reaches from the cardiac region to the cloaca. The right and left kidneys are parallel to each other, quite close together, and only separated by the mesentery--the thin narrow layer that attaches the middle gut to the under surface of the vertebral column. The passage of each primitive kidney, the nephroduct, runs towards the back on the lower and outer side of the gland, and opens in the cloaca, close to the starting-point of the allantois; it afterwards opens into the allantois itself.

(FIGURES 2.396 TO 2.398. Urinary and sexual organs of ox-embryos. Figure 2.396, female embryo one and a half inches long; Figure 2.397, male embryo, one and a half inches long. Figure 2.398 female embryo two and a half inches long. w primitive kidney, wg Wolffian duct, m Mullerian duct, m apostrophe upper end of same (opened at t), i lower and thicker part of same (rudiment of uterus), g genital cord, h testicle, (h apostrophe, lower and h double apostrophe, upper testicular ligament), o ovary, o apostrophe lower ovarian ligament, i inguinal ligament of primitive kidney, d diaphragmatic ligament of primitive kidney, nn accessory kidneys, n permanent kidneys, under them the S-shaped ureters, between these the rectum, v bladder, a umbilical artery. (From Kolliker.))

The primitive or primordial kidneys of the amniote embryo were formerly called the "Wolffian bodies," and sometimes "Oken's bodies." They act for a time as kidneys, absorbing unusable juices from the embryonic body and conducting them to the cloaca--afterwards to the allantois. There the primitive urine accumulates, and thus the allantois acts as bladder or urinary sac in the embryos of man and the other Amniotes. It has, however, no genetic connection with the primitive kidneys, but is a pouch-like growth from the anterior wall of the rectum (Figure 1.147 u). Thus it is a product of the visceral layer, whereas the primitive kidneys are a product of the middle layer. Phylogenetically we must suppose that the allantois originated as a pouch-like growth from the cloaca-wall in consequence of the expansion caused by the urine accumulated in it and excreted by the kidneys. It is originally a blind sac of the rectum. The real bladder of the vertebrate certainly made its first appearance among the Dipneusts (in Lepidosiren), and has been transmitted from them to the Amphibia, and from these to the Amniotes. In the embryo of the latter it protrudes far out of the not yet closed ventral wall. It is true that many of the fishes also have a "bladder." But this is merely a local enlargement of the lower section of the nephroducts, and so totally different in origin and composition from the real bladder. The two structures can be compared from the physiological point of view, and so are ANALOGOUS, as they have the same function; but not from the morphological point of view, and are therefore not HOMOLOGOUS. The false bladder of the fishes is a mesodermic product of the nephroducts; the true bladder of the Dipneusts, Amphibia, and Amniotes is an entodermic blind sac of the rectum.

In all the Anamnia (the lower amnionless Craniotes, Cyclostomes, Fishes, Dipneusts, and Amphibia) the urinary organs remain at a lower stage of development to this extent, that the primitive kidneys (protonephri) act permanently as urinary glands. This is only so as a passing phase of the early embryonic life in the three higher classes of Vertebrates, the Amniotes. In these the permanent or after or secondary (really tertiary) kidneys (renes or metanephri) that are distinctive of these three classes soon make their appearance. They represent the third and last generation of the vertebrate kidneys. The permanent kidneys do not arise (as was long supposed) as independent glands from the alimentary tube, but from the last section of the primitive kidneys and the nephroduct. Here a simple tube, the secondary renal duct, develops, near the point of its entry into the cloaca; and this tube grows considerably forward. With its blind upper or anterior end is connected a glandular renal growth, that owes its origin to a differentiation of the last part of the primitive kidneys. This rudiment of the permanent kidneys consists of coiled urinary canals with Malpighian capsules and vascular coils (without ciliated funnels), of the same structure as the segmental mesonephridia of the primitive kidneys. The further growth of these metanephridia gives rise to the compact permanent kidneys, which have the familiar bean-shape in man and most of the higher mammals, but consist of a number of separate folds in the lower mammals, birds, and reptiles. As the permanent kidneys grow rapidly and advance forward, their passage, the ureter, detaches altogether from its birth-place, the posterior end of the nephroduct; it passes to the posterior surface of the allantois. At first in the oldest Amniotes this ureter opens into the cloaca together with the last section of the nephroduct, but afterwards separately from this, and finally into the permanent bladder apart from the rectum altogether. The bladder originates from the hindmost and lowest part of the allantoic pedicle (urachus), which enlarges in spindle shape before the entry into the cloaca. The anterior or upper part of the pedicle, which runs to the navel in the ventral wall of the embryo, atrophies subsequently, and only a useless string-like relic of it is left as a rudimentary organ; that is the single vesico-umbilical ligament. To the right and left of it in the adult male are a couple of other rudimentary organs, the lateral vesico-umbilical ligaments. These are the degenerate string-like relics of the earlier umbilical arteries.

Though in man and all the other Amniotes the primitive kidneys are thus early replaced by the permanent kidneys, and these alone then act as urinary organs, all the parts of the former are by no means lost. The nephroducts become very important physiologically by being converted into the passages of the sexual glands. In all the Gnathostomes--or all the Vertebrates from the fishes up to man--a second similar canal develops beside the nephroduct at an early stage of embryonic evolution. The latter is usually called the Mullerian duct, after its discoverer, Johannes Muller, while the former is called the Wolffian duct. The origin of the Mullerian duct is still obscure; comparative anatomy and ontogeny seem to indicate that it originates by differentiation from the Wolffian duct. Perhaps it would be best to say: "The original primary nephroduct divides by differentiation (or longitudinal cleavage) into two secondary nephroducts, the Wolffian and the Mullerian ducts." The latter (Figure 2.387 m) lies just on the inner side of the former (Figure 2.387 w). Both open behind into the cloaca.

However uncertain the origin of the nephroduct and its two products, the Mullerian and the Wolffian ducts, may be, its later development is clear enough. In all the Gnathostomes the Wolffian duct is converted into the spermaduct, and the Mullerian duct into the oviduct. Only one of them is retained in each sex; the other either disappears altogether, or only leaves relics in the shape of rudimentary organs. In the male sex, in which the two Wolffian ducts become the spermaducts, we often find traces of the Mullerian ducts, which I have called "Rathke's canals" (Figure 2.394 c). In the female sex, in which the two Mullerian ducts form the oviducts, there are relics of the Wolffian ducts, which are called "the ducts of Gaertner."

(FIGURE 2.399. Female sexual organs of a Monotreme (Ornithorhynchus, Figure 2.269). o ovaries, t oviducts, u womb, sug urogenital sinus; at u apostrophe is the outlet of the two wombs, and between them the bladder (vu). cl cloaca. (From Gegenbaur.)

FIGURES 2.400 AND 2.401. Original position of the sexual glands in the ventral cavity of the human embryo (three months old).

FIGURE 2.400 male (natural size). h testicles, gh conducting ligament of the testicles, wg spermaduct, h bladder, uh inferior vena cava, nn accessory kidneys, n kidneys.

FIGURE 2.401 female, slightly magnified. r round maternal ligament (underneath it the bladder, over it the ovaries). r apostrophe kidneys, s accessory kidneys, c caecum, o small reticle, om large reticle (stomach between the two), l spleen. (From Kolliker.))

We obtain the most interesting information with regard to this remarkable evolution of the nephroducts and their association with the sexual glands from the Amphibia (Figures 2.390 to 2.395). The first structure of the nephroduct and its differentiation into Mullerian and Wolffian ducts are just the same in both sexes in the Amphibia, as in the mammal embryos (Figures 2.392 and 2.396). In the female Amphibia the Mullerian duct develops on either side into a large oviduct (Figure 2.393 od), while the Wolffian duct acts permanently as ureter (u). In the male Amphibia the Mullerian duct only remains as a rudimentary organ without any functional significance, as Rathke's canal (Figure 2.394 c); the Wolffian duct serves also as ureter, but at the same time as spermaduct, the sperm-canals (ve) that proceed from the testicles (t) entering the fore part of the primitive kidneys and combining there with the urinary canals.

In the mammals these permanent amphibian features are only seen as brief phases of the earlier period of embryonic development (Figure 2.392). Here the primitive kidneys, which act as excretory organs of urine throughout life in the amnion-less Vertebrates, are replaced in the mammals by the permanent kidneys. The real primitive kidneys disappear for the most part at an early stage of development, and only small relics of them remain. In the male mammal the epididymis develops from the uppermost part of the primitive kidney; in the female a useless rudimentary organ, the epovarium, is formed from the same part. The atrophied relic of the former is known as the paradidymis, that of the latter as the parovarium.

(FIGURE 2.402. Urogenital system of a human embryo of three inches in length, double natural size. h testicles, wg spermaducts, gh conducting ligament, p processus vaginalis, b bladder, au umbilical arteries, m mesorchium, d intestine, u ureter, n kidney, nn accessory kidney. (From Kollman.))

The Mullerian ducts undergo very important changes in the female mammal. The oviducts proper are developed only from their upper part; the lower part dilates into a spindle-shaped tube with thick muscular wall, in which the impregnated ovum develops into the embryo. This is the womb (uterus). At first the two wombs (Figure 2.399 u) are completely separate, and open into the cloaca on either side of the bladder (vu), as is still the case in the lowest living mammals, the Monotremes. But in the Marsupials a communication is opened between the two Mullerian ducts, and in the Placentals they combine below with the rudimentary Wolffian ducts to form a single "genital cord." The original independence of the two wombs and the vaginal canals formed from their lower ends are retained in many of the lower Placentals, but in the higher they gradually blend and form a single organ. The conjunction proceeds from below (or behind) upwards (or forwards). In many of the Rodents (such as the rabbit and squirrel) two separate wombs still open into the simple and single vaginal canal; but in others, and in the Carnivora, Cetacea, and Ungulates, the lower halves of the wombs have already fused into a single piece, though the upper halves (or "horns") are still separate ("two-horned" womb, uteris bicornis). In the bats and lemurs the "horns" are very short, and the lower common part is longer. Finally, in the apes and in man the blending of the two halves is complete, and there is only the one simple, pear-shaped uterine pouch, into which the oviducts open on each side. This simple uterus is a late evolutionary product, and is found ONLY in the ape and man.

(FIGURES 2.403 TO 2.406. Origin of human ova in the female ovary.

FIGURE 2.403. Vertical section of the ovary of a new-born female infant, a ovarian epithelium, b rudimentary string of ova, c young ova in the epithelium, d long string of ova with follicle-formation (Pfluger's tube), e group of young follicles, f isolated young follicle, g blood-vessels in connective tissue (stroma) of the ovary. In the strings the young ova are distinguished by their considerable size from the surrounding follicle-cells. (From Waldeyer.)

FIGURE 2.404. Two young Graafian follicles, isolated. In 1 the follicle-cells still form a simple, and in 2 a double, stratum round the young ovum; in 2 they are beginning to form the ovolemma or the zona pellucida (a).

FIGURES 2.405 AND 2.406. Two older Graafian follicles, in which fluid is beginning to accumulate inside the eccentrically thickened epithelial mass of the follicle-cells (Figure 2.405 with little, 2.406 with much, follicle-water). ei the young ovum, with embryonic vesicle and spot, zp ovolemma or zona pellucida, dp discus proligerus, formed of an accumulation of follicle-cells, which surround the ovum, ff follicle-liquid (liquor folliculi), gathered inside the stratified follicle-epithelium (fe), fk connective-tissue fibrous capsule of the Graafian follicle (theca folliculi).)

In the male mammals there is the same fusion of the Mullerian and Wolffian ducts at their lower ends. Here again they form a single genital cord (Figure 2.397 g), and this opens similarly into the original urogenital sinus, which develops from the lowest section of the bladder (v). But while in the male mammal the Wolffian ducts develop into the permanent spermaducts, there are only rudimentary relics left of the Mullerian ducts. The most notable of these is the "male womb" (uterus masculinus), which originates from the lowest fused part of the ducts, and corresponds to the female uterus. It is a small, flask-shaped vesicle without any physiological significance, which opens into the ureter between the two spermaducts and the prostate folds (vesicula prostatica).

(FIGURE 2.407. A ripe human Graafian follicle. a the mature ovum, b the surrounding follicle-cells, c the epithelial cells of the follicle, d the fibrous membrane of the follicle, e its outer surface.)

The internal sexual organs of the mammals undergo very distinctive changes of position. At first the germinal glands of both sexes lie deep inside the ventral cavity, at the inner edge of the primitive kidneys (Figures 2.386 g and 2.392 k), attached to the vertebral column by a short mesentery (mesorchium in the male, mesovarium in the female). But this primary arrangement is retained permanently only in the Monotremes (and the lower Vertebrates). In all other mammals (both Marsupials and Placentals) they leave their original cradle and travel more or less far down (or behind), following the direction of a ligament that goes from the primitive kidneys to the inguinal region of the ventral wall. This is the inguinal ligament of the primitive kidneys, known in the male as the Hunterian ligament (Figure 2.400 gh), and in the female as the "round maternal ligament" (Figure 2.401 r). In woman the ovaries travel more or less towards the small pelvis, or enter into it altogether. In the male the testicles pass out of the ventral cavity, and penetrate by the inguinal canal into a sac-shaped fold of the outer skin. When the right and left folds ("sexual swellings") join together they form the scrotum. The various mammals bring before us the successive stages of this displacement. In the elephant and the whale the testicles descend very little, and remain underneath the kidneys. In many of the rodents and carnassia they enter the inguinal canal. In most of the higher mammals they pass through this into the scrotum. As a rule, the inguinal canal closes up. When it remains open the testicles may periodically pass into the scrotum, and withdraw into the ventral cavity again in time of rut (as in many of the marsupials, rodents, bats, etc.).

The structure of the external sexual organs, the copulative organs that convey the fecundating sperm from the male to the female organism in the act of copulation, is also peculiar to the mammals. There are no organs of this character in most of the other Vertebrates. In those that live in water (such as the Acrania and Cyclostomes, and most of the fishes) the ova and sperm-cells are simply ejected into the water, where their conjunction and fertilisation are left to chance. But in many of the fishes and amphibia, which are viviparous, there is a direct conveyance of the male sperm into the female body; and this is the case with all the Amniotes (reptiles, birds, and mammals). In these the urinary and sexual organs always open originally into the last section of the rectum, which thus forms a cloaca (Chapter 2.22). Among the mammals this arrangement is permanent only in the Monotremes, which take their name from it (Figure 2.399 cl). In all the other mammals a frontal partition is developed in the cloaca (in the human embryo about the beginning of the third month), and this divides it into two cavities. The anterior cavity receives the urogenital canal, and is the sole outlet of the urine and the sexual products; the hind or anus-cavity passes the excrements only.

Even before this partition has been formed in the Marsupials and Placentals, we see the first trace of the external sexual organs. First a conical protuberance rises at the anterior border of the cloaca-outlet--the sexual prominence (phallus, Figure 2.402 A, e, B, e). At the tip it is swollen in the shape of a club ("acorn" glans). On its under side there is a furrow, the sexual groove (sulcus genitalis, f), and on each side of this a fold of skin, the "sexual pad" (torus genitalis, h l). The sexual protuberance or phallus is the chief organ of the sexual sense (Chapter 2.25); the sexual nerves spread on it, and these are the principal organs of the specific sexual sensation. As erectile bodies (corpora cavernosa) are developed in the male phallus by peculiar modifications of the blood-vessels, it becomes capable of erecting periodically on a strong accession of blood, becoming stiff, so as to penetrate into the female vagina and thus effect copulation. In the male the phallus becomes the penis; in the female it becomes the much smaller clitoris; this is only found to be very large in certain apes (Ateles). A prepuce ("foreskin") is developed in both sexes as a protecting fold on the anterior surface of the phallus.

(FIGURE 408. The human ovum after issuing from the Graafian follicle, surrounded by the clinging cells of the discus proligerus (in two radiating crowns). z ovolemma (zona pellucida, with radial porous canals), p cytosoma (protoplasm of the cell-body, darker within, lighter without), k nucleus of the ovum (embryonic vesicle). (From Nagel, magnified 250 times.) (Cf. Figures 1.1 and 1.14.)

The external sexual member (phallus) is found at various stages of development within the mammal class, both in regard to size and shape, and the differentiation and structure of its various parts; this applies especially to the terminal part of the phallus, the glans, both the larger glans penis of the male and the smaller glans clitoridis of the female. The part of the cloaca from the upper wall of which it forms belongs to the proctodaeum, the ectodermic invagination of the rectum (Chapter 2.27); hence its epithelial covering can develop the same horny growths as the corneous layer of the epidermis. Thus the glans, which is quite smooth in man and the higher apes, is covered with spines in many of the lower apes and in the cat, and in many of the rodents with hairs (marmot) or scales (guinea-pig) or solid horny warts (beaver). Many of the Ungulates have a free conical projection on the glans, and in many of the Ruminants this "phallus-tentacle" grows into a long cone, bent hook-wise at the base (as in the goat, antelope, gazelle, etc.). The different forms of the phallus are connected with variations in the structure and distribution of the sensory corpuscles--i.e. the real organs of the sexual sense, which develop in certain papillae of the corium of the phallus, and have been evolved from ordinary tactile corpuscles of the corium by erotic adaptation (Chapter 2.25).

The formation of the corpora cavernosa, which cause the stiffness of the phallus and its capability of penetrating the vagina, by certain special structures of their spongy vascular spaces, also shows a good deal of variety within the vertebrate stem. This stiffness is increased in many orders of mammals (especially the carnassia and rodents) by the ossification of a part of the fibrous body (corpus fibrosum). This penis-bone (os priapi) is very large in the badger and dog, and bent like a hook in the marten; it is also very large in some of the lower apes, and protrudes far out into the glans. It is wanting in most of the anthropoid apes; it seems to have been lost in their case (and in man) by atrophy.

The sexual groove on the under side of the phallus receives in the male the mouth of the urogenital canal, and is changed into a continuation of this, becoming a closed canal by the juncture of its parallel edges, the male urethra. In the female this only takes place in a few cases (some of the lemurs, rodents, and moles); as a rule, the groove remains open, and the borders of this "vestibule of the vagina" develop into the smaller labia (nymphae). The large labia of the female develop from the sexual pads (tori genitales), the two parallel folds of the skin that are found on each side of the genital groove. They join together in the male, and form the closed scrotum. These striking differences between the two sexes cannot yet be detected in the human embryo of the ninth week. We begin to trace them in the tenth week of development, and they are accentuated in proportion as the difference of the sexes develops.

Sometimes the normal juncture of the two sexual pads in the male fails to take place, and the sexual groove may also remain open (hypospadia). In these cases the external male genitals resemble the female, and they are often wrongly regarded as cases of hermaphrodism. Other malformations of various kinds are not infrequently found in the human external sexual organs, and some of them have a great morphological interest. The reverse of hypospadia, in which the penis is split open below, is seen in epispadia, in which the urethra is open above. In this case the urogenital canal opens above at the dorsal root of the penis; in the former case down below. These and similar obstructions interfere with a man's generative power, and thus prejudicially affect his whole development. They clearly prove that our history is not guided by a "kind Providence," but left to the play of blind chance.

We must carefully distinguish the rarer cases of real hermaphrodism from the preceding. This is only found when the essential organs of reproduction, the genital glands of both kinds, are united in one individual. In these cases either an ovary is developed on the right and a testicle on the left (or vice versa); or else there are testicles and ovaries on both sides, some more and others less developed. As hermaphrodism was probably the original arrangement in all the Vertebrates, and the division of the sexes only followed by later differentiation of this, these curious cases offer no theoretical difficulty. But they are rarely found in man and the higher mammals. On the other hand, we constantly find the original hermaphrodism in some of the lower Vertebrates, such as the Myxinoides, many fishes of the perch-type (serranus), and some of the Amphibia (ringed snake, toad). In these cases the male often has a rudimentary ovary at the fore end of the testicle; and the female sometimes has a rudimentary, inactive testicle. In the carp also and some other fishes this is found occasionally. We have already seen how traces of the earlier hemaphrodism can be traced in the passages of the Amphibia.

Man has faithfully preserved the main features of his stem-history in the ontogeny of his urinary and sexual organs. We can follow their development step by step in the human embryo in the same advancing gradation that is presented to us by the comparison of the urogenital organs in the Acrania, Cyclostomes; Fishes, Amphibia, Reptiles, and then (within the mammal series) in the Monotremes, Marsupials, and the various Placentals. All the peculiarities of urogenital structure that distinguish the mammals from the rest of the Vertebrates are found in man; and in all special structural features he resembles the apes, particularly the anthropoid apes. In proof of the fact that the special features of the mammals have been inherited by man, I will, in conclusion, point out the identical way in which the ova are formed in the ovary. In all the mammals the mature ova are contained in special capsules, which are known as the Graafian follicles, after their discoverer, Roger de Graaf (1677). They were formerly supposed to be the ova themselves; but Baer discovered the ova within the follicles (Chapter 1.3). Each follicle (Figure 2.407) consists of a round fibrous capsule (d), which contains fluid and is lined with several strata of cells (c). The layer is thickened like a knob at one point (b); this ovum-capsule encloses the ovum proper (a). The mammal ovary is originally a very simple oval body (Figure 2.387 g), formed only of connective tissue and blood-vessels, covered with a layer of cells, the ovarian epithelium or the female germ epithelium. From this germ epithelium strings of cells grow out into the connective tissue or "stroma" of the ovary (Figure 2.403 b). Some of the cells of these strings (or Pfluger's tubes) grow larger and become ova (primitive ova, c); but the great majority remain small, and form a protective and nutritive stratum of cells round each ovum--the "follicle-epithelium" (e).

The follicle-epithelium of the mammal has at first one stratum (Figure 2.404 1), but afterwards several (2). It is true that in all the other Vertebrates the ova are enclosed in a membrane, or "follicle," that consists of smaller cells. But it is only in the mammals that fluid accumulates between the growing follicle-cells, and distends the follicle into a large round capsule, on the inside wall of which the ovum lies, at one side (Figures 2.405 and 2.406). There again, as in the whole of his morphology, man proves indubitably his descent from the mammals.

In the lower Vertebrates the formation of ova in the germ-epithelium of the ovary continues throughout life; but in the higher it is restricted to the earlier stages, or even to the period of embryonic development. In man it seems to cease in the first year; in the second year we find no new-formed ova or chains of ova (Pfluger's tubes). However, the number of ova in the two ovaries is very large in the young girl; there are calculated to be 72,000 in the sexually-mature maiden. In the production of the ova men resemble most of the anthropoid apes.

Generally speaking, the natural history of the human sexual organs is one of those parts of anthropology that furnish the most convincing proofs of the animal origin of the human race. Any man who is acquainted with the facts and impartially weighs them will conclude from them alone that we have been evolved from the lower Vertebrates. The larger and the detailed structure, the action, and the embryological development of the sexual organs are just the same in man as in the apes. This applies equally to the male and the female, the internal and the external organs. The differences we find in this respect between man and the anthropoid apes are much slighter than the differences between the various species of apes. But all the apes have certainly a common origin, and have been evolved from a long-extinct early-Tertiary stem-form, which we must trace to a branch of the lemurs. If we had this unknown pithecoid stem-form before us, we should certainly put it in the order of the true apes in the primate system; but within this order we cannot, for the anatomic and ontogenetic reasons we have seen, separate man from the group of the anthropoid apes. Here again, therefore, on the ground of the pithecometra-principle, comparative anatomy and ontogeny teach with full confidence the descent of man from the ape.