perforated by small apertures. Most internal of all is a layer of endothelial cells, which form the free surface over which the blood flows. The arteries are not nourished by the blood which flows through them, but by minute vessels, vasa vasorum, distributed in their external, elastic and muscular coats.
|Fig. 2.—Diagram of the structure of an artery. A, tunica adventitia;|
E, elastic coat; M, muscular coat; F, fenestrated coat;
En, endothelium continuous with the endothelial wall of
C, the capillaries.
The earliest appearance of the blood vessels is dealt with under Vascular System. Here will be briefly described the fate of the main vessel which carries the blood away from the truncus arteriosus of the developing heart (q.v.). This ventral aorta, if traced forward, soon divides into two lateral parts, the explanation being that there were originally two vessels, side by side, which fused to form the heart, but continued separate anteriorly. The two parts run for a little distance toward the head of the embryo, ventral to the alimentary canal, and then turn toward the dorsum, passing one on either side of that tube to form the first aortic arch. Having reached the dorsum they turn backward toward the tail end and form the dorsal aortae; here, according to A. H. Young (Studies in Anatomy, Owens College, 1891 and 1900) they again turn toward the ventral side and become, after a transitional stage, the hypogastric, placental, allantoic or umbilical arteries. This authority does not believe that the middle sacral artery of the adult is the real continuation of the single median dorsal aorta into which the two parallel dorsal vessels just mentioned soon coalesce, though until recently it has always been so regarded. The anterior loop between the ventral and dorsal aortae already described as the first aortic arch is included in the maxillary or first visceral arch of the soft parts (see fig. 3, 1). Later, four other well-marked aortic arches grow behind this in the more caudal visceral arches, so that there are altogether five arterial arches on each side of the pharynx, through which the blood can pass from the ventral to the dorsal aorta. Of these arches the first soon disappears, but is probably partly represented in the adult by the internal maxillary artery, one branch of which, the infraorbital, is enclosed in the upper jaw, while another, the inferior dental, is surrounded by the lower jaw. Possibly the ophthalmic artery also belongs to this arch. The second arch also disappears, but the posterior auricular and occipital arteries probably spring from it, and at an early period it passed through the stapes as the transitory stapedial artery. The third arch forms the beginning of the internal carotid. The fourth arch becomes the arch of the adult aorta, between the origins of the left carotid and left subclavian, on the left side, and the first part of the right subclavian artery on the right. The apparent fifth arch on the left side (fig. 3, 6) remains all through foetal life as the ductus arteriosus, and, as the lungs develop, the pulmonary arteries are derived from it. J. E. V. Boas and W. Zimmermann have shown that this arch is in reality the sixth, and that there is a very transitory true fifth arch in front of it (fig. 3, 5). The part of the ventral aorta from which this last arch rises is a single median vessel due to the same fusion of the two primitive ventral aortae which precedes the formation of the heart, but a spiral septum has appeared in it which divides it in such a way that while the anterior or cephalic arches communicate with the left ventricle of the heart, the last one communicates with the right (see Heart). The fate of the ventral and dorsal longitudinal vessels must now be followed. The fused part of the two ventral aortae, just in front of the heart, forms the ascending part of the adult aortic arch, and where this trunk divides between the fifth and fourth arches (strictly speaking, the sixth and fifth), the right one forms the innominate (fig. 3, In.) and the left one a very short part of the transverse arch of the aorta until the fourth arch comes off (see fig. 4). From this point to the origin of the third arch is common carotid, and after that, to the head, external carotid on each side. The dorsal longitudinal arteries on the head side of the junction with the third arch form the internal carotids. Between the third and fourth arches they are obliterated, while on the caudal side of this, until the point of fusion is reached on the dorsal side of the heart, the left artery forms the upper part of the dorsal aorta while the right entirely disappears. Below this point the thoracic and abdominal aortae are formed by the two primitive dorsal aortae which have fused to form a single median vessel. As the limbs are developed, vessels bud out in them. The subclavian for the arm comes from the fourth aortic arch on each side, while in the leg the main artery is a branch of the caudal arch which is curving ventralward to form the umbilical artery. From the convexity of this arch the internal iliac and sciatic at first carry the blood to the limb, as they do permanently in reptiles, but later the external iliac and femoral become developed, and, as they are on the concave side of the bend of the hip, while the sciatic is on the convex, they have a mechanical advantage and become the permanent main channel.
For further details see O. Hertwig, Handbuch der vergleichenden und experimentellen Entwickelungslehre der Wirbeltiere (Jena, 1905).
In the Acrania the lancelet (Amphioxus) shows certain arrangements of its arteries which are suggestive of the embryonic stages of the higher vertebrates and Man. There is a median ventral aorta below the pharynx, from which branchial arteries run up on each side between the branchial clefts, where the blood is aerated, to join two dorsal aortae which run back side by side until the hind end of the pharynx is reached; here they fuse to form a median vessel from which branches are distributed to the straight intestine. There is no heart, but the ventral aorta is contractile, and the blood is driven forward in it and backward in the dorsal aortae. The branchial arteries are very numerous, and cannot be homologized closely with the five (originally six) pairs of aortic arches in Man.
In the fish the ventral aorta gives rise to five afferent branchial arteries carrying the blood to the gills, though these may not all come off as independent trunks from the aorta. From the gills the afferent branchials carry the blood to the median dorsal aorta. As pectoral and pelvic fins are now developed, subclavian and iliac arteries are found rising from the dorsal aorta, though the aorta itself is continued directly backward as the caudal artery into the tail. In the Dipnoi or mud fish, in which the swim bladder is converted into a functional lung, the hindmost afferent branchial artery, corresponding to the fifth (strictly speaking the sixth) aortic arch of the human embryo, gives off on each side a pulmonary artery to that structure.The arrangement of the branchial aortic arches in the tailed Amphibia (Urodela), and in the tadpole stage of the tailless forms (Anura), makes it probable that the generalized vertebrate has six (if not more) pairs of these instead of the five which are evident in the human embryo. Four pairs of arches are present, the first of which is the carotid and corresponds to the third of Man; the second is the true aortic arch on each side; the third undergoes