The first constraint on vascular development is physiological. Unlike new machines, which do not need to function until they leave the assembly line, new organisms have to function even as they develop. Embryonic cells must obtain nourishment before there is an intestine, use oxygen before there are lungs, and excrete wastes before there are kidneys. All these functions are mediated through the embryonic circulatory system. Therefore, the circulatory physiology of the developing embryo must differ from that of the adult organism. Food is absorbed not through the intestine, but from either the yolk or the placenta, and respiration is conducted not through the gills or lungs, but through the chorionic or allantoic membranes (see Figure 12.1). The major embryonic blood vessels must be constructed to serve these extraembryonic structures.

The second constraint is evolutionary. The mammalian embryo extends blood vessels to the yolk sac even though there is no yolk inside. Moreover, the blood leaving the heart via the truncus arteriosus passes through vessels that loop over the foregut to reach the dorsal aorta. Six pairs of these aortic arches loop over the pharynx (Figure 1). In primitive fish, these arches persist and enable the gills to oxygenate the blood through the gills. In adult birds and mammals, in which lungs oxygenate the blood, such a system makes little sense—but all six pairs of aortic arches are formed in mammalian and avian embryos before the system eventually becomes simplified into a single aortic arch. Thus, even though our physiology does not require such a structure, our embryonic condition reflects our evolutionary history.

Figure 1  Aortic arches of the human embryo. (A) Originally, the truncus arteriosus pumps blood into the aorta, which branches on either side of the foregut. The six aortic arches take blood from the truncus arteriosus and allow it to flow into the dorsal aorta. (B) As development proceeds, arches begin to disintegrate or become modified (the dotted lines indicate degenerating structures). The first two pairs of arches completely disappear. The third arches become the internal carotid arteries. The right fourth arch becomes the right subclavian artery, and the left fourth arch becomes the major arch of the aorta. The fifth arches disintegrate. The right sixth arch disappears, but the left sixth arch gives rise to the pulmonary arteries and the ductus arteriosus (which will start closing on the first breath). (C) Eventually, the remaining arches are modified and the adult arterial system is formed. However, numerous variations of this scheme are found in the human population.

The third set of constraints is physical. According to the laws of fluid movement, the most effective transport of fluids is performed by large tubes. As the radius of a blood vessel gets smaller, resistance to flow increases as r–4 (Poiseuille’s law). A blood vessel that is half as wide as another has a resistance to flow 16 times greater. However, diffusion of nutrients can take place only when blood flows slowly and has access to cell membranes. So here is a paradox: the constraints of diffusion mandate that vessels be small, whereas the laws of hydraulics mandate that vessels be large. This paradox has been solved by the evolution of circulatory systems with a hierarchy of vessel sizes (LaBarbera 1990). In dogs, for example, blood in the large vessels (aorta and vena cava) flows more than 100 times faster than it does in the capillaries. With a system of large vessels specialized for transport and small vessels specialized for diffusion (where the blood spends most of its time), nutrients and oxygen can reach the individual cells of the growing organism. This hierarchy is seen very early in development; it is already well established in the 3-day chick embryo.

But this is not the entire story. If fluid under constant pressure moves directly from a large-diameter tube into a small-diameter tube (as in a hose nozzle), the fluid velocity increases. The evolutionary solution to this problem was the emergence of many smaller vessels branching out from a larger one, making the collective cross-sectional area of all the smaller vessels greater than that of the larger vessel. Circulatory systems show a relationship (known as Murray’s law) in which the cube of the radius of the parent vessel approximates the sum of the cubes of the radii of the smaller vessels. Computer models of blood vessel formation must take into account not only gene expression patterns but also the fluid dynamics of blood flow (Gödde and Kurz 2001). The construction of any circulatory system negotiates among all of these physical, physiological, and evolutionary constraints.




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