The alligator and its allies

Body flexure has increased until now the forebrain and tail are almost in contact (Fig. 17). The eye has developed somewhat; the ear vesicle, which is not shown in the figure, is small and seems to lie nearer the ventral side; the nasal pit is much larger and is crescentic in shape. The hyomandibular cleft (g′) still persists as a small crescentic slit, while the next three clefts are now represented merely by superficial grooves separated by distinct ridges, the visceral folds. No indication of a fifth cleft is seen. The maxillary process (mx) grows ventralward under the forebrain and is already longer than the mandibular arch (md).

The chief advance in development over the preceding stage, besides the formation of the maxillary process, is in the appearance of the appendages (aa and pa); they have the characteristic shape of the rudimentary vertebrate appendage, though the anterior pair seem to point in an unusual direction at this stage and to be slightly more developed than the posterior. The curious, anteriorly directed heart (ht) is, perhaps, somewhat abnormal. The umbilical stalk (u) is comparatively narrow and, like the allantois, was cut off close to the body.

Transverse sections of an embryo of this stage are represented in Figures 17a-g, drawn under a lower magnification than were any of the preceding figures.

Figure 17a is in the region of the pharynx, and passes through the forebrain (fb) and posterior part of the hindbrain (hb). In the thick walls of both of these structures histological differentiation has begun, so that even under low power an inner granular and an outer clear zone may be distinguished. Under greater magnification the presence of short fibers may be made out among the cells. The cerebral hemispheres (ch) are well-marked structures, their asymmetry being of course due to the obliquity of the section. Only one eye is cut by the plane of the section, and this one shows no connection with the forebrain. The outer wall of the optic cup (oc) is so thin that under this magnification it can scarcely be seen as a dark line surrounding the retinal wall. The lens (ln) is now a solid mass, of the usual type for vertebrate embryos, its front or outer wall being a scarcely discernible line. The hindbrain (hb) has the usual form for that region and does not differ particularly from what was noted in earlier stages except in the histological differentiation that has already been mentioned. As with the eye, it is only on the right side that the auditory vesicle (o) is shown. It shows some differentiation, but not so much as would be seen were it cut in another region. In the center of the section the pharynx (ph) forms an irregular cavity connected with the exterior on the left by a gill cleft (g) and by another slit which is simply the anterior margin of the stomodæum. On the right neither of these openings is in the plane of the figure, though the gill cleft (hyomandibular), which lies close to the auditory vesicle, is almost an open passage. A few small blood-vessels are scattered through the section; one of these (bv), lying between the notochord (nt) and the floor of the brain, is noticeable from its being very closely packed with corpuscles, so that at first glance, under low magnification, it looks more like a nerve than a blood-vessel.

Figure 17b is also through the pharyngeal region, a short distance behind the preceding section. The growth of the cerebral hemispheres (ch) is better shown than in the preceding figure, as is also the general form of the optic cup (oc). On the left the nasal cavity (n) is seen as an elongated slit with thick walls; it is cut near, but not through, its opening to the exterior. The same gill cleft (g) that was seen in the preceding figure is seen here as a narrow, transverse cleft, open at both ends. Between the notochord (nt) and the spinal cord (sc) is the same, though now double, blood-filled vessel (bv) that was seen in the preceding section. The other blood-vessels are larger here than in the more anterior region. There is a faint condensation of mesoblast in the neighborhood of the notochord, and a more marked condensation (mp) farther toward each side is the curiously shaped muscle plate.

Figure 17c is through the heart region, and that organ is cut through the opening from the lower or ventricular into the upper or auricular chamber. The thickening of the wall of the ventricle, which was noticed in the preceding stage, has increased to such an extent that there is now a marked difference in the thickness of the ventricular and auricular walls. As in the preceding stage, the body wall is torn, probably in handling, so that it appears to be incomplete around the ventral side of the heart. Dorsal to the heart two small circular holes (ent) with thick walls are the œsophagus and trachea, cut anterior to the point of bifurcation of the latter into the bronchial or lung rudiments. On either side of these structures is an elongated blood-vessel (ac), the anterior cardinal vein, its elongation being due to the fact that it is cut at the place where it turns downward to empty into the heart. Dorsal to the œsophagus are the aortæ (ao), which are here cut just at the point where the two vessels unite to form one; the next section, posterior to the one under discussion, shows an unpaired aorta. The notochord (nt) and spinal cord (sc) need no description, except to note that the latter shows active histological differentiation, numerous mitotic figures being seen under higher magnification, especially in the cells that line the spinal canal. On the right of the cord the edge of a spinal ganglion (sg) is seen, in connection with which in other sections are seen the clearly defined nerve roots. The condensation of mesoblast around the notochord is quite evident, and in close contact with this medial condensation are two very characteristic, S-shaped muscle plates (mp), which extend from the level of the dorsal side of the spinal cord to the upper limits of the cardinal veins. In some sections the muscle plates even yet show slight remains of the myocœl at the dorsal end.

Figure 17d is in the region of the posterior end of the heart (ht), which is cut through the tip of the ventricle, and the anterior end of the liver (li), which has the appearance of a mass of darkly stained cords or strands of cells surrounding a large blood-vessel (mv). This blood-vessel may be called the meatus venosus, though it is not separated by any line of demarcation from the auricle. A few sections anterior to this region the meatus venosus opens dorsally into a large vessel on each side (dc), which at first glance seems a part of the body cavity, but which is in reality the ductus Cuvieri, formed by the union of the anterior and posterior cardinal veins. An irregular, crescentic cleft (bc), lying medial and parallel to each of the Cuvierian vessels, is the body cavity. In the upper angle of this cavity is a granular mass, the glomerulus, that of the left side being accompanied by the extreme anterior end of the Wolffian duct. In the rounded mass of mesoblast, between the cleft-like regions of the body cavity, the lung rudiments (lu), and the œsophagus (oe) are seen as three small, circular openings; that of the œsophagus is somewhat smaller than the other two. The notochord (nt), spinal cord (sc), and muscle plates (mp) have almost the same appearance as in the preceding section. A spinal ganglion (sg) is seen on each side of the spinal cord; the one on the left shows a well-defined spinal nerve (sn), which may be traced ventrally as far as the end of the muscle plate, along whose medial side it courses. The ventral nerve root is plainly seen; the dorsal root, in this section, less plainly. The amnion (a) and abdominal wall are, as in the preceding figure, torn in the region of the ventricle.

Figure 17e is a short distance posterior to the figure just described. The liver is cut through its middle region and forms a large, darkly staining, reticular mass on the left side of the figure. The digestive tract is seen at two places to the right of the liver; the smaller and more ventral of these openings (i) may be called the intestine, while the larger is evidently the stomach (i′). The body wall is here unfused and becomes suddenly thinner as it passes upward into the amnion (a). The Wolffian tubules (wt) form a very conspicuous mass on either side of the mesentery, in close connection with the posterior cardinal veins (pc). In the mesoblast between the dorsal aorta (ao) and the notochord are two small, irregular, darkly stained masses (sy). These are shown in the preceding two figures, but were not mentioned in the description. They may be traced through a great part of the length of the embryo back of the head region; at intervals corresponding in length to the distance between the spinal ganglia they are enlarged, while between these enlargements they are very small in cross-section. At certain points a small blood-vessel is given off by the dorsal aorta to the immediate neighborhood of each of these small areas. Although they show no connection with the central nervous system, these structures appear to be the rudiments of the sympathetic nerves.

Figure 17f is in the region just in front of the hind legs. The abdominal walls are here unfused, and into the unenclosed body cavity projects the intestine (i), supported by a narrow mesentery and surrounded by a comparatively thick mass of mesoblast. The Wolffian body and duct form a mass of considerable size on each side of the mesentery. The Wolffian body is cut near its posterior end and consists of smaller tubules than in the more anterior regions. The Wolffian ducts (wd), on the other hand, are very large and are much more clearly distinguishable from the Wolffian tubules than in the preceding sections. The Wolffian ridges (wr) are very marked projections on the sides of the body, and in a region farther caudad become especially developed as the posterior appendages, to be described in connection with the following section. Both spinal ganglia are shown in this figure (sg), and in connection with the left ganglion the spinal nerve (sn), extending ventrally as far as the level of the Wolffian duct. The sympathetic nerve rudiments do not extend so far caudad as the plane of this section. The dorsal end of each muscle plate (mp) is seen, in this and other sections, to be slightly enlarged to form a round knob; this knob contains a distinct cavity (not shown in the figure), the myocœl.

In Figure 17g, owing to the curvature of the body, the plane of the section passes through the body at three places: through the region of the heart and lungs (Fig. 17d), through the region of the posterior appendages, and through the tail. In fact, the plane of the section represented by each of the preceding figures cut the embryo in more than one region, but for the sake of simplicity only one region was represented in each figure. In the figure under discussion only the leg and tail regions have been drawn, though the latter region (t), being cut through one of its curves, is seen as an elongated body with a section of the spinal cord, notochord, etc., at each end. Both regions shown in the figure are enclosed in the same fold (a) of the amnion. Of the structures in the dorsal side of the larger or more anterior part of this figure nothing need be said. The most striking feature of the section is the presence of the large posterior leg rudiments (pa). As was noted in the preceding figure, they are, as usual, merely local enlargements or projections of the mesoblast (covered, of course, with ectoblast) of the Wolffian ridge. They are, as shown in this section and in the surface view of this stage (Fig. 17), bluntly pointed projections from the sides of the body. The anterior appendage seems to be slightly more developed than the posterior, as was noted in describing the surface view of the embryo. The digestive tract is cut through its extreme posterior end, in the region that may be termed the cloaca (cl), for into it at this point the Wolffian ducts open (wdo). As the narrow cloacal chamber is followed toward the tail, it becomes still smaller in diameter, and the ventral depression or cleft seen in this figure gradually becomes deeper until its walls are continuous with the ectoderm that covers the ventral projection of mesoderm between the hind legs; no actual opening to the exterior is present, however. There is a space of about twenty-five or thirty sections (in a series of eight hundred) between the posterior ends of the Wolffian bodies and the cloacal openings of the Wolffian ducts. The body cavity (bc) and the posterior cardinal veins (pc) are very small in this region, as might be expected.

STAGE XV

Figure 18 (Plate XXIV.)

Only the head of this embryo is represented, as the general state of development is about the same as in the preceding stage.

The chief object in making the figure is to show the five gill clefts (g¹⁻⁵). The fifth cleft, though small and probably not open to the exterior, is quite distinct in this embryo. The writer would feel more doubt of its being a true, though rudimentary, gill cleft had not Clarke (17) found a fifth pair of clefts in his material. The nasal pit has advanced in development somewhat and shows the beginning of the groove that connects it with the mouth. In this figure the crescentic hyomandibular cleft shows its connection with the groove between the mandibular and the hyoid folds.

STAGE XVI

Figure 19 (Plate XXIV.)

This embryo is only slightly more developed than the preceding. Body flexure is so great that the forebrain and tail nearly touch. Only the anterior three gill clefts are visible. The maxillary process (mx) is longer and more narrow; the mandibular fold has not changed appreciably. The nasal pit (n) is now connected by a distinct groove with the stomodæum. The appendages have increased in size, the posterior (pa) being the longer. The anterior appendage (aa) is distinctly broadened to form the manus, while no sign of the pes is to be seen at the extremity of the posterior appendage. The heart (ht) is still very prominent. The stalk of the umbilicus (u), which is quite narrow, projects from the ventral wall in the region between the heart and the hind legs. The tail is of considerable length and is closely coiled.

STAGE XVII

Figures 20-20j (Plates XXV., XXVI.)

The superficial changes noted in this stage chiefly concern the head, which has increased considerably in length (Fig. 20). The curvature of the body is slightly more marked, and the tail is more tightly coiled at the end. There are still signs of three gill clefts. The maxillary process (mx) is long and narrow, while the mandibular arch (md) is still short and broad. The fronto-nasal region has greatly increased and has the aquiline profile noted by Clarke. The nasal groove has disappeared, and there remains the small opening (n) at the side of the fronto-nasal region, near the end of the still separate maxillary process. The umbilicus is in about the same condition as in the preceding stage, but the heart is less prominent. The outline of the manus (ma) is more definite, and the extremity of the posterior appendage is distinctly flattened out to form the rudimentary pes (pe). The position of the elbow-joint in the anterior appendage is seen at the end of the reference line aa.

Typical transverse sections of this stage are shown in Figures 20a-j.

Figure 20a is a section through the middle region of the head, cutting the hindbrain on one side and the forebrain on the other. The walls of the brain show rather more histological differentiation than was seen in the preceding sections, though this cannot be shown under the low magnification used. The hindbrain (hb), which is cut near its anterior border, exhibits the usual membranous dorsal and thick ventral walls. The forebrain is here seen as three distinct cavities—a median third ventricle (tv), with a thick ventral wall, and a thin dorsal wall extended to form a large paraphysis (epi), and two lateral ventricles (ch), the cavities of the cerebral hemispheres, whose walls are quite thick except on the side next the third ventricle. The sections of this series being slightly oblique, the eye is here cut on the right side only, where it is seen as a large, semicircular cavity (e) with thick, dense walls. The mesoblast, in which several blood-vessels (bv) are seen, exhibits three distinct areas—a median, lighter zone, with a more dense area on either side. The significance of this variation in the density of the mesoblast is not apparent.

Figure 20b is only a few sections posterior to the section just described. It is drawn chiefly to show the appearance of the forebrain, the other structures being about as in the preceding figure, except that both eyes (e) are here represented. The section passes through the wide opening between the third (tv) and the lateral ventricles (ch) and cuts the anterior edge of the pineal body[8] (epi). The paraphysis is very large and is directed backward instead of forward, as is usually the case among the lower vertebrates (if the alligator may be so classed). It is shown in Figure 17a of a preceding stage and will be again shown in a sagittal section to be described later. The same areas of more dense and less dense mesoblast noted in the preceding figure are seen here.

Figure 20c, though still in the head region, shows several features that were not seen in the preceding figures. On the left of the hindbrain (hb) the auditory vesicle (o), which is now considerably more advanced than in earlier figures, is seen as a larger, flask-shaped cavity and a smaller, round one. Between the larger cavity and the hindbrain is the root of a cranial nerve (cn), apparently the eighth, since in another section it comes in close contact with the wall of the larger part of the auditory vesicle just mentioned. On the right side, ventral to the hindbrain, another and much larger nerve (cn) is seen. Nearly in the center of the figure is seen a small, irregular, thick-walled cavity (p); this is the pituitary body, and its connection with the roof of the pharynx may easily be made out in another section. The mesoblast in this region of the sections contains numerous large and small blood-vessels and exhibits certain denser areas which probably represent the beginnings of the cranial cartilages. No sign of the forebrain is seen (the plane of the section passing in front of that region), except that the tip of the wall of one of the cerebral hemispheres (ch) is cut. The left nasal chamber (n) is shown: it will be noted again in the following section. The eye on the right side shows no remarkable features; its lens (ln) is large and lies well back of the lips of the optic cup, which may now be called the iris (ir). A thin layer of mesoblast has pushed in between the lens and the superficial ectoderm to form the cornea, and the outer wall of the optic cup is now distinctly pigmented. The inner wall of the optic cup is beginning to differentiate into the retinal elements. The eye on the left side is cut farther from its central region and has a very different appearance from the eye just described. This unusual appearance is due to the fact that the section passed through the choroid fissure, which is very large and seems to be formed by the pushing in of the walls of the cup and not by a mere cleft in these walls. This fissure is hardly noticeable in the stage preceding the present, and in a stage slightly older it has disappeared; so that it would seem to be a very transient structure. It apparently is formed at about the time that the optic stalk, as such, disappears. It is in the region of the choroid fissure, if not through it, that the optic nerve (on) enters the eye. Through the fissure also enters a vascular tuft of mesoblast (pt) which may be seen projecting into the optic cup after the disappearance of the fissure. This loop of blood-vessels is doubtless the pecten.

Figure 20d represents a section through the hindbrain (hb), pharynx (ph), and tip of the snout. On either side of the hindbrain are a convoluted auditory vesicle (o), and several blood-vessels and nerves, while ventral to it is seen the anterior end of the notochord (nt), around which the mesoblast is somewhat more dense than elsewhere. The pharynx (ph) sends out toward the surface a narrow gill cleft (g′) in the neighborhood of each auditory vesicle. These clefts connect with the exterior by very narrow slits, not seen in the plane of this section. The opposite end of the pharynx, as seen in this figure, opens on the left (pn) into the nasal chamber. The nasal cavity on the right is cut in such a plane that it shows neither its external nor its pharyngeal opening. The nasal passages are here fairly long and nearly straight chambers; their lining epithelium is quite thick in the middle region, but becomes thinner where it merges into the epithelium of the pharynx at one end, and into the superficial epithelium at the other end. The unusual appearance of the eye (e), on the right side of the figure, is due to the fact that the plane of the section cut tangentially through the extreme edge of the eye in the region of the choroid fissure.

Figure 20e is only a short distance posterior to the preceding. On the left side the pharynx (ph) is connected with the exterior through the stomodæum, and on the right the hyomandibular cleft (g′) is cut almost through its opening to the exterior. The auditory vesicle (o) on the right is cut near its middle region, while that on the left is barely touched by the plane of the section. The notochord (nt), with its condensed area of mesoblast, is somewhat larger than in the preceding section. The nasal canal on the right (n) is cut through neither anterior nor posterior opening, while on the left side the canal shows the anterior opening (an).

Figure 20f, which is in the region of the posterior part of the pharynx and the anterior part of the heart, shows some rather unusual conditions.

The spinal cord (sc) and notochord (nt), with the faintly outlined condensations of mesoblast in their region, need no special description. The pharynx (ph) is here reduced to an irregular, transversely elongated cavity, the lateral angles of which are connected on each side with the exterior through a tortuous and almost closed gill cleft (g), which must be followed through many sections before its inner and outer openings may be determined. Dorsal to the pharynx numerous blood-vessels (bv), both large and small, may be seen, while ventral to it is noticed a faint condensation of mesoblast (la), in the form of an inverted T, the anlage of the laryngeal structures. The ventral portion of the figure is made up of a nearly circular thin-walled cavity, the pericardium (pr). Most of the pericardial cavity is occupied in this section by the thick-walled ventricle (vn), above which is the bulbus (b) and the tip of the auricle (au). The bulbus is nearly circular in outline, though its cavity is very irregular. A few sections anterior to this, the opening of the bulbus into the ventricle is seen.

In Figure 20g the section represented is only a short distance posterior to the one represented by Figure 20f. The mesoblastic structures in the neighborhood of the spinal cord (sc) and notochord (nt) will be described in connection with the next figure, where they are more clearly defined. The œsophagus (oe)—or posterior end of the pharynx, whichever it may be called—is here a crescentic slit, with its convex side upward; ventrally it opens by a narrow glottis into the trachea (ta). The trachea is surrounded by the same condensed area of mesoblast (la) that was mentioned in connection with the preceding figure, but the condensation is here more marked. From the bulbus (b) an aortic arch (ar) extends upward for a short distance on the right side, while to the left of the œsophagus an aortic arch (ar) is cut through the upper part of its course. Ventral to the bulbus the ventricle (vn) and two auricles (au) are seen surrounded by the pericardial wall.

Figure 20h is in the region of the liver (li), which has about the same position in relation to the auricles (au) that was occupied by the ventricle in the last figure. The auricles are connected with each other by a wide passage. The trachea (ta) and the œsophagus (oe) are entirely distinct from each other; the former is a small, nearly circular hole, while the lumen of the latter is obliterated and its walls form a solid, bow-shaped mass of cells. Since there is a narrow space between this mass of cells and the surrounding mesoblast, it might be thought that the lumen of the œsophagus had been closed by the simple shrinkage of its walls; higher magnification, however, fails to show any sign of a collapsed lumen. It is doubtless the problematic and temporary closure of the œsophagus that is noticed in other forms. On each side of the œsophagus, in close relation with the anterior cardinal vein (ac), is noticed a nerve (cn) cut through a ganglionic enlargement. When traced forward these nerves are seen to arise from the region of the medulla, and when followed caudad they are found to be distributed chiefly to the tissues surrounding the newly formed bronchi; they are doubtless the tenth cranial nerves. On the right side of the figure the close connection of this nerve with the near-by gill cleft is seen. Above the paired aortæ (ao) the sympathetic nerves (sy) will be noticed. The mesoblast surrounding the spinal cord (sc) and notochord (nt) is distinctly condensed (more so than the figure shows) to form what may be called the centrum (c) and neural arch (na) of the vertebræ. The arch, owing to the slight obliquity of the section, shows here only on one side. The spinal cord is not yet completely enclosed by the neural arches. The muscle plates (mp) are in close connection with the rudiments of the vertebræ just mentioned. The spinal cord (sc) is here differentiated into three areas—a dense, deeply stained area immediately around the neurocœl; a less dense area of cells surrounding the inner area and extending ventralward as a rounded projection on each side; and an outer layer, with few or on nuclei, surrounding the inner two layers except on the dorsal side.

In Figure 20i the size and complexity of the figure are due, it will be easily understood, to the fact that the plane of the section passed through the curve of the body, thus practically cutting the embryo in two regions—an anterior, where the lungs (lu) and liver (li) are seen, and a posterior, where the Wolffian bodies (wt) are present. The spinal cord and the surrounding structures have almost the same characteristics at both ends of the figure, except that the primitive spinal column is rather more distinct in the posterior end of the section. The posterior cardinal veins (pc), Wolffian ducts (wd), and Wolffian bodies (wt) are also prominent structures of this end of the figure, the last being made up of a great number of tubules. The extreme anterior ends of the Wolffian bodies are seen in the other half of the section in the upper angles of the body cavity, dorsal to the lung rudiments (lu). Filling most of the body cavity (bc) and making up the greater part of the middle of the figure are the liver (li), now a very large organ; the stomach (i′), also quite large; the pancreas (pan), a small body lying near the stomach; and the lungs (lu), which here consist of several thick-walled tubes, surrounded by lobes of mesoblast. The other features of the figure need no special mention.

Figure 20j is through the base of the posterior appendages (pa), in which the cartilages are already being outlined by condensations of mesoblast. The intestine (i) is cut in two regions—at a more anterior point, where it is seen as a small, circular hole surrounded by mesoblast and hung by a narrow mesentery, and through the cloacal region, the larger and more ventral cavity, into which the Wolffian ducts (wd) open a short distance caudad to this section. The blood-vessels present a rather curious appearance. A short distance anterior to this point the aorta has divided into three, or it might be said that it has given off two, large branches. These two branches, one on either side near the posterior cardinal vein, pass toward the ventral side of the embryo on each side of the cloaca and end at about the region represented by the present figure. The small portion of the aorta that remains after the giving off of the two branches just described continues, as the caudal artery (ca), into the tail; it is a small vessel just under the notochord, and gives off small, paired branches at regular intervals toward the vertebral region. The posterior cardinal veins (pc), posterior to the openings of the Wolffian ducts into the cloaca, unite to form a large caudal vein lying just ventral to the caudal artery.

STAGE XVIII

Figure 21 (Plate XXVII.)

This embryo, as may be seen, for example, by the form of the appendages, is slightly further developed than the one represented in Figure 20. The figure is from a photograph of a living embryo as it lay in the egg, a portion of the shell and shell membranes having been removed. The embryo, which lies on its left side, is rather faintly outlined because of the overlying allantois. The allantois has been increasing rapidly in size, and is here so large that it extends beneath the cut edges of the shell at all points except in the region in front of the head of the embryo, where its border may be seen. Its blood-vessels, especially the one that crosses the head just back of the eye, are clearly shown in the figure, and in the living specimen, when filled with the bright red blood, they form a most beautiful demonstration. As in the chick, the allantois lies close beneath the shell membranes and is easily torn in removing them.

STAGE XIX

Figure 22 (Plate XXVII.)

Figure 22 is a photograph of a somewhat older embryo, removed from the egg and freed of the fetal membranes. The appendages show the position of both elbow and knee joints, and in the paddle-shaped manus and pes the digits may be faintly seen. The tail is very long and is spirally coiled, the outer spiral being in contact with the frontal region of the head. The jaws are completely formed, the upper projecting far beyond the lower. The elliptical outline of the eyes is noticeable, but the lids are still too little developed to be seen in this figure. The surface of the embryo is still smooth and white.

STAGE XX

Figures 23-23b (Plate XXVII.)

In this surface view (Fig. 23) several changes are seen, though no very great advance in development has taken place. The outlines of the digits (five in the manus and four in the pes) are now well de fined; they even project slightly beyond the general outline of the paddle-shaped part. The tail has begun to straighten out, and it now extends across the front of the face. The lower jaw has increased in length, but is still shorter than the upper. The eyelids, especially the upper, are beginning to be discernible in surface view. Though still without pigment, the surface of the body is beginning to show by faint transverse lines the development of scales; these lines are most evident in this figure in the middle region of the tail, just before it crosses the nose.

A sagittal section of the entire embryo (except the tail) of this age is shown in Figure 23a. In the head region the section is nearly median, while the posterior part of the body is cut slightly to one side of the middle line. At the tip of the now well-developed snout is seen one of the nostrils (an), cut through the edge; its connection with the complicated nasal chamber (n) is not here seen, nor is the connection of the nasal chamber with the posterior nares (pn). The pharynx (ph), is anteriorly connected with the exterior through the mouth (m) and the nares, while posteriorly it opens into the œsophagus (oe); the trachea (ta), though distinct from the œsophagus, does not yet open into the pharynx. In the lower jaw two masses of cartilage are seen, one near the symphysis (mk) and one near the wall of the trachea, doubtless the rudiment of the hyoid. The deep groove back of the Meckel’s cartilage (mk) marks the tip of the developing tongue, which here forms the thick mass on the floor of the mouth cavity. Dorsal to the pharynx a mass of cartilage (se) is developing in the sphenethmoid region. This being a median section, the ventricles of the fore- (fb), mid- (mb), and hindbrain (hb) are seen as large cavities, while the cerebral hemispheres (ch) appear nearly solid, only a small portion of a lateral ventricle showing. The paraphysis (epi) is cut a little to one side of the middle and so does not show its connection with the brain. At the base of the brain the infundibulum (in) is seen as an elongated cavity whose ventral wall is in close contact with a group of small, darkly staining alveoli (p), the pituitary body. Extending posteriorly from the pituitary body is a gradually thickening mass of cartilage (bp), which surrounds the anterior end of the notochord (nt) and may be called the basilar plate. In its anterior region, where the section is nearly median, the spinal column shows its canal, with the enclosed spinal cord, while toward the posterior end of the figure the vertebræ are cut to one side of the middle line, and hence show the neural arches (na) with the alternating spinal ganglia (sg). Near the posterior end of the figure the pelvic girdle (pl) is seen. The largest organ of the embryo, as seen in this section, is the heart, of which the ventricle (vn) seems to be closely surrounded, both in front and behind, by the auricles (au). The liver (li) is the large, reticular mass back of the heart. Dorsal and anterior to the liver is the lung (lu), now of considerable size and development. The enteron is cut in several places (oe, i) and its walls are beginning to show some differentiation, though this cannot be seen under the magnification here used. One of the Wolffian bodies is seen as a huge mass of tubules (wt) extending from the pelvic region, where the mass is greatest, to the region of the lungs. The Wolffian tubules stain darkly and the whole structure forms a very striking feature of the section. Dorsal to the posterior end of the Wolffian body is a small, oval mass of very fine tubules (k), which do not stain so darkly as do the Wolffian tubules; this mass is apparently the beginning of the permanent kidney, the metanephros. Its tubules, though their origin has not been determined, seem to be entirely distinct from the tubules of the Wolffian body.

A single vertical section through the anterior part of the head of an embryo of this age has been represented in Figure 23b. On the right side the plane of the section cut through the lens of the eye (ln); on the left side the section was anterior to the lens. The upper (ul) and lower (ll) eyelids are more evident here than in the surface view. Owing to the hardness of the lens, its supporting structures were torn away in sectioning. The vitreous humor is not represented in the figure. The superior (ur) and inferior (lr) recti muscles are well shown on the right side; they are attached to the median part of a Y-shaped mass of cartilage (se), which may be termed the sphenethmoidal cartilage. Between the branches of this Y-shaped cartilage the anterior ends of the cerebral hemispheres (ch)—better called, perhaps, the olfactory lobes—are seen. Between the lower end of the sphenethmoidal cartilage and a dorsally evaginated part of the pharynx are two small openings (pn); when traced forward these tubes are found to open into the convoluted nasal chamber, while a short distance posterior to the plane of this figure they unite with each other and open almost immediately into the pharynx. The rather complicated structures of the nasal passages of the alligator have been described by the writer in another paper (57). In the lower jaw the cartilage (mk) is seen on either side and several bands of muscle are developing in the mesoblast. Two deep grooves give form to what may be called the rudimentary tongue (tn). In both jaws one or two tooth rudiments (to) may be distinguished as small invaginations of ectoderm.

STAGE XXI

Figure 24 (Plate XXVII.)

In this stage the curvature of the body and tail is less marked than was seen in the last surface view. The body has increased greatly in size, so that the size of the head is relatively not so great. The size of the eye in relation to that of the head is much diminished also. The five anterior and four posterior digits are well formed, and their claws are of considerable size, though of course not present on all the digits. The outlines of scales may be traced from the tip of the tail to the skull; they are especially prominent along the dorsal profile. The skin is just beginning to show traces of pigment, which is, however, not shown in the photograph. The umbilical stalk is seen projecting with a loop of the intestine from the abdominal wall; this is shown more clearly in the next stage. The embryo now begins to exhibit some of the external characteristics of the adult alligator.

STAGE XXII

Figure 25 (Plate XXVIII.)

This embryo needs no particular description. It has reached in its external appearance practically the adult condition, although there is still considerable yolk (not shown in the figure) to be absorbed, and the embryo would not have hatched for many days. Pigmentation, begun in the last stage, is now complete. The umbilical stalk is clearly seen projecting from a large opening in the body wall. The long loop of the intestine that extends down into the yolk sac is here evident, and it is hard to understand how it can all be drawn up into the body cavity when the umbilical stalk is withdrawn. No sharp shell-tooth at the tip of the snout, such as is described by Voeltzkow (78) in the crocodile, is here seen.

STAGE XXIII

Figure 26 (Plate XXVIII.)

This figure shows the relative sizes of the just-hatched alligator and the egg from which it came. It also shows the position of the young alligator in the egg, half of the shell having been removed for that purpose. The blotchy appearance of the unopened egg is due chiefly to stains produced by the decayed vegetation of the nest. At hatching the young alligator is about 20 cm. long, nearly three times the length of the egg; but the tail is so compressed that, though it makes up about half of the length of the animal, it takes up very little room in the egg.

SUMMARY

Owing to the fact that the embryo may undergo considerable development before the egg is laid, and also to the unusual difficulty of removing the very young embryos, the earlier stages of development are very difficult to obtain.

The mesoderm seems to be derived chiefly by proliferation from the entoderm, in which way all of that anterior to the blastopore arises. Posterior to the blastopore the mesoderm is proliferated from the lower side of the ectoderm in the usual way. No distinction can be made between the mesoderm derived from the ectoderm and that derived from the entoderm.

The ectoderm shows during the earlier stages a very great increase in thickness along the median longitudinal axis of the embryo.

The notochord is apparently of entodermal origin, though in the posterior regions, where the germ layers are continuous with each other, it is difficult to decide with certainty.

The medullary folds have a curious origin, difficult to explain without the use of figures. They are continuous posteriorly with the primitive streak, so that it is impossible to tell where the medullary groove ends and the primitive groove begins, unless the dorsal opening of the blastopore be taken as the dividing point.

The amnion develops rapidly, and entirely from the anterior end.

The blastopore or neurenteric canal is a very distinct feature of all the earlier stages up to about the time of closure of the medullary canal.

Preceding the ordinary cranial flexure there is a sort of temporary bending of the head region, due apparently to the formation of the head-fold.

During the earlier stages of development the anterior end of the embryo is pushed under the surface of the blastoderm, and is hence not seen from above.

Body torsion is not so definite in direction as in the chick, some embryos lying on the right side, others on the left.

Of the gill clefts, three clearly open to the exterior and probably a fourth also. A probable fifth cleft was seen in sections and in one surface view.

The first trace of the urinary system is seen as a dorsally projecting, solid ridge of mesoblast in the middle region of the embryo, which ridge soon becomes hollowed out to form the Wolffian duct.

The origin of the hypophysis and paraphysis is clearly seen; the latter projects backward.

No connection can be seen between the first rudiments of the sympathetic nerves and the central nervous system.

The lumen of the œsophagus is for a time obliterated as in other forms.

The choroid fissure is a very transitory but well-marked feature of the eye.

LETTERING FOR ALL FIGURES ON PLATES VI.-XXVIII.

• a, head-fold of amnion.
• aa, anterior appendage.
• ac, anterior cardinal vein.
• al, allantois.
• an, anterior nares.
• ao, aorta.
• aop, area opaca.
• ap, area pellucida.
• ar, aortic arch.
• au, auricle.
• b, bulbus arteriosus.
• bc, body cavity.
• blp, blastopore.
• bp, basilar plate.
• bv, blood-vessel.
• c, centrum of vertebra.
• ca, caudal artery.
• ch, cerebral hemisphere.
• cl, cloaca.
• cn, cranial nerve.
• cp, posterior choroid plexus.
• cv, cardinal veins.
• dc, ductus Cuvieri.
• e, eye.
• ec, ectoderm.
• ec′, thickening of ectoderm,
• en, entoderm.
• en′, endocardium.
• ent, enteron.
• ep, epidermal layer of ectoderm.
• epi, paraphysis.
• es, embryonic shield.
• f, fronto-nasal process.
• fb, forebrain.
• fg, foregut.
• g¹⁻⁵, gill clefts.
• gf¹⁻⁶, gill folds.
• gl, glomerulus.
• h, head-fold.
• hb, hindbrain.
• ht, heart.
• i, intestine.
• i′, stomach.
• in, infundibulum.
• ir, iris.
• it, iter.
• k, kidney (metanephros).
• l, remains of groove between secondary folds.
• la, larynx (cartilages of).
• li, liver.
• ll, lower lid of eye.
• ln, lens.
• lr, inferior rectus muscle of eye.
• lu, lungs.
• lv, lens vesicle.
• m, mouth.
• ma, manus.
• mb, midbrain.
• mc, medullary canal.
• me′, tip end of medullary canal.
• md, mandibular fold.
• mes, mesoderm.
• mes′, myocardium.
• mf, medullary fold.
• mg, medullary groove.
• mk, Meckel’s cartilage.
• mp, muscle plate.
• ms, mesentery.
• mv, meatus venosus.
• mx, maxillary fold.
• myc, myocœl.
• n, nasal invagination or cavity.
• na, neural arch of vertebra.
• nc, neurenteric canal.
• nl, nervous layer of ectoderm.
• nt, notochord.
• o, ear vesicle.
• oc, optic cup.
• oe, œsophagus.
• on, optic nerve.
• os, optic stalk.
• ov, optic vesicle.
• p, pituitary body.
• pa, posterior appendage.
• pan, pancreas.
• pc, posterior cardinal vein.
• pe, pes.
• pg, primitive groove.
• ph, pharynx.
• pl, pelvis.
• pn, posterior nares.
• pr, pericardial cavity.
• ps, primitive streak.
• pt, pecten.
• rt, retina.
• s, somites.
• sc, spinal cord.
• se, sphenethmoid cartilage.
• sf, secondary fold.
• sg, spinal ganglion.
• sm, splanchnic mesoblast.
• sn, spinal nerve.
• so, somatic mesoblast.
• st, stomodæum.
• sy, sympathetic nervous system.
• t, tail.
• ta, trachea.
• tg, thyroid gland.
• th, thickening and posterior limit of sf.
• tn, tongue.
• to, tooth anlage.
• tr, truncus arteriosus.
• tv, third ventricle of brain.
• tv′, third ventricle of brain.
• u, umbilical stalk.
• ul, upper lid of eye.
• ur, superior rectus muscle of eye.
• v′-″-‴, first, second, and third cerebral vesicles.
• va, vascular area.
• vm, vitelline membrane.
• vn, ventricle of heart.
• vv, vitelline blood-vessels.
• wd, Wolffian duct.
• wdo, opening of Wolffian duct.
• wr, Wolffian ridge.
• wt, Wolffian tubules.
• y, yolk.

EXPLANATION OF FIGURES 1-26 ON PLATES VI.-XXVIII.