human cardiovascular system

human cardiovascular system


      organ system that conveys blood through vessels to and from all parts of the body, carrying nutrients and oxygen to tissues and removing carbon dioxide and other wastes. It is a closed tubular system in which the blood is propelled by a muscular heart. Two circuits, the pulmonary and the systemic, consist of arterial (artery), capillary, and venous (vein) components.

      The primary function of the heart is to serve as a muscular pump propelling blood into and through vessels (blood vessel) to and from all parts of the body. The arteries, which receive this blood at high pressure and velocity and conduct it throughout the body, have thick walls that are composed of elastic fibrous tissue and muscle cells. The arterial tree—the branching system of arteries—terminates in short, narrow, muscular vessels called arterioles, from which blood enters simple endothelial tubes (i.e., tubes formed of endothelial, or lining, cells) known as capillaries. These thin, microscopic capillaries are permeable to vital cellular nutrients and waste products that they receive and distribute. From the capillaries, the blood, now depleted of oxygen and burdened with waste products, moving more slowly and under low pressure, enters small vessels called venules that converge to form veins, ultimately guiding the blood on its way back to the heart.

      This article describes the structure and function of the heart and blood vessels, and the technologies that are used to evaluate and monitor the health of these fundamental components of the human cardiovascular system. For a discussion of diseases affecting the heart and blood vessels, see the article cardiovascular disease. For a full treatment of the composition and physiologic function of blood, see blood, and for more information on diseases of the blood, see blood disease. To learn more about the human circulatory system, see systemic circulation and pulmonary circulation, and for more about cardiovascular and circulatory function in other living organisms, see circulation.

The heart

Shape and location
 The human adult heart is normally slightly larger than a clenched fist with average dimensions of about 13 × 9 × 6 centimetres (5 × 3.5 × 2.5 inches) and weighing approximately 10.5 ounces (300 grams). It is cone-shaped, with the broad base directed upward and to the right and the apex pointing downward and to the left. It is located in the chest (thoracic (thoracic cavity)) cavity behind the breastbone ( sternum), in front of the windpipe ( trachea), the esophagus, and the descending aorta, between the lungs, and above the diaphragm (the muscular partition between the chest and abdominal cavities). About two-thirds of the heart lies to the left of the midline.

      The heart is suspended in its own membranous sac, the pericardium. The strong outer portion of the sac, or fibrous pericardium, is firmly attached to the diaphragm below, the mediastinal pleura on the side, and the sternum in front. It gradually blends with the coverings of the superior vena cava and the pulmonary (lung) arteries and veins (pulmonary circulation) leading to and from the heart. (The space between the lungs, the mediastinum, is bordered by the mediastinal pleura, a continuation of the membrane lining the chest. The superior vena cava is the principal channel for venous blood from the chest, arms, neck, and head.)

      Smooth, serous (moisture-exuding) membrane lines the fibrous pericardium, then bends back and covers the heart. The portion of membrane lining the fibrous pericardium is known as the parietal serous layer (parietal pericardium), that covering the heart as the visceral serous layer (visceral pericardium or epicardium).

      The two layers of serous membrane are normally separated only by 10 to 15 millilitres (0.6 to 0.9 cubic inch) of pericardial fluid, which is secreted by the serous membranes. The slight space created by the separation is called the pericardial cavity. The pericardial fluid lubricates the two membranes with every beat of the heart as their surfaces glide over each other. Fluid is filtered into the pericardial space through both the visceral and parietal pericardia.

Chambers of the heart
      The heart is divided by septa, or partitions, into right and left halves, and each half is subdivided into two chambers. The upper chambers, the atria (atrium), are separated by a partition known as the interatrial septum; the lower chambers, the ventricles (ventricle), are separated by the interventricular septum. The atria receive blood from various parts of the body and pass it into the ventricles. The ventricles, in turn, pump blood to the lungs and to the remainder of the body.

      The right atrium, or right superior portion of the heart, is a thin-walled chamber receiving blood from all tissues except the lungs. Three veins empty into the right atrium, the superior and inferior venae cavae, bringing blood from the upper and lower portions of the body, respectively, and the coronary sinus, draining blood from the heart itself. Blood flows from the right atrium to the right ventricle. The right ventricle, the right inferior portion of the heart, is the chamber from which the pulmonary artery carries blood to the lungs.

      The left atrium, the left superior portion of the heart, is slightly smaller than the right atrium and has a thicker wall. The left atrium receives the four pulmonary veins, which bring oxygenated blood from the lungs. Blood flows from the left atrium into the left ventricle. The left ventricle, the left inferior portion of the heart, has walls three times as thick as those of the right ventricle. Blood is forced from this chamber through the aorta to all parts of the body except the lungs.

External surface of the heart
      Shallow grooves called the interventricular sulci, containing blood vessels, mark the separation between ventricles on the front and back surfaces of the heart. There are two grooves on the external surface of the heart. One, the atrioventricular groove, is along the line where the right atrium and the right ventricle meet; it contains a branch of the right coronary artery (the coronary arteries deliver blood to the heart muscle). The other, the anterior interventricular sulcus, runs along the line between the right and left ventricles and contains a branch of the left coronary artery.

      On the posterior side of the heart surface, a groove called the posterior longitudinal sulcus marks the division between the right and left ventricles; it contains another branch of a coronary artery. A fourth groove, between the left atrium and ventricle, holds the coronary sinus, a channel for venous blood.

Origin and development
      In the embryo, formation of the heart begins in the pharyngeal, or throat, region. The first visible indication of the embryonic heart occurs in the undifferentiated mesoderm, the middle of the three primary layers in the embryo, as a thickening of invading cells. An endocardial (lining) tube of flattened cells subsequently forms and continues to differentiate until a young tube with forked anterior and posterior ends arises. As differentiation and growth progress, this primitive tube begins to fold upon itself, and constrictions along its length produce four primary chambers. These are called, from posterior to anterior, the sinus venosus, atrium, ventricle, and truncus arteriosus. The characteristic bending of the tube causes the ventricle to swing first to the right and then behind the atrium, the truncus coming to lie between the sideways dilations of the atrium. It is during this stage of development and growth that the first pulsations (pulse) of heart activity begin.

      Endocardial cushions (local thickenings of the endocardium, or heart lining) “pinch” the single opening between the atrium and the ventricle into two portions, thereby forming two openings. These cushions are also responsible for the formation of the two atrioventricular valves (the valves between atria and ventricles), which regulate the direction of blood flow through the heart.

      The atrium becomes separated into right and left halves first by a primary partition with a perforation and later by a secondary partition, which, too, has a large opening, called the foramen ovale, in its lower part. Even though the two openings do not quite coincide in position, blood still passes through, from the right atrium to the left. At birth, increased blood pressure in the left atrium forces the primary partition against the secondary one, so that the two openings are blocked and the atria are completely separated. The two partitions eventually fuse.

      The ventricle becomes partially divided into two chambers by an indentation of myocardium (heart muscle) at its tip. This developing partition is largely muscular and is supplemented by membranous connective tissue that develops in conjunction with the subdivision of the truncus arteriosus by a spiral partition into two channels, one for systemic and one for pulmonary circulation (the aorta and the pulmonary artery, respectively). At this time, the heart rotates clockwise and to the left so that it resides in the left thorax, with the left chambers posterior and the right chambers anterior. The greater portion of blood passing through the right side of the heart in the fetus is returned to the systemic circulation by the ductus arteriosus, a vessel connecting the pulmonary artery and the aorta. At birth this duct becomes closed by a violent contraction of its muscular wall. Thereafter the blood in the right side of the heart is driven through the pulmonary arteries to the lungs for oxygenation and returned to the left side of the heart for ejection into the systemic circulation. A distinct median furrow at the apex of the ventricles marks the external subdivision of the ventricle into right and left chambers.

Structure and function
Valves (valve) of the heart
      To prevent backflow of blood, the heart is equipped with valves that permit the blood to flow in only one direction. There are two types of valves located in the heart: the atrioventricular valves (tricuspid and mitral) and the semilunar valves (pulmonary and aortic).

      The atrioventricular valves are thin, leaflike structures located between the atria and the ventricles. The right atrioventricular opening is guarded by the tricuspid valve, so called because it consists of three irregularly shaped cusps, or flaps. The leaflets consist essentially of folds of endocardium (the membrane lining the heart) reinforced with a flat sheet of dense connective tissue. At the base of the leaflets, the middle supporting flat plate becomes continuous with that of the dense connective tissue of the ridge surrounding the openings.

      Tendinous cords of dense tissue (chordae tendineae) covered by thin endocardium extend from the nipplelike papillary muscles to connect with the ventricular surface of the middle supporting layer of each leaflet. The chordae tendineae and the papillary muscles from which they arise limit the extent to which the portions of the valves near their free margin can billow toward the atria. The left atrioventricular opening is guarded by the mitral, or bicuspid, valve, so named because it consists of two flaps. The mitral valve is attached in the same manner as the tricuspid, but it is stronger and thicker because the left ventricle is by nature a more powerful pump working under high pressure.

      Blood is propelled through the tricuspid and mitral valves as the atria contract. When the ventricles contract, blood is forced backward, passing between the flaps and walls of the ventricles. The flaps are thus pushed upward until they meet and unite, forming a complete partition between the atria and the ventricles. The expanded flaps of the valves are restrained by the chordae tendineae and papillary muscles from opening into the atria.

      The semilunar valves are pocketlike structures attached at the point at which the pulmonary artery and the aorta leave the ventricles. The pulmonary valve guards the orifice between the right ventricle and the pulmonary artery. The aortic valve protects the orifice between the left ventricle and the aorta. The three leaflets of the aortic semilunar and two leaflets of the pulmonary valves are thinner than those of the atrioventricular valves, but they are of the same general construction with the exception that they possess no chordae tendineae.

      Closure of the heart valves is associated with an audible sound, called the heartbeat. The first sound occurs when the mitral and tricuspid valves close, the second when the pulmonary and aortic semilunar valves close. These characteristic heart sounds have been found to be caused by the vibration of the walls of the heart and major vessels around the heart. The low-frequency first heart sound is heard when the ventricles contract, causing a sudden backflow of blood that closes the valves and causes them to bulge back. The elasticity of the valves then causes the blood to bounce backward into each respective ventricle. This effect sets the walls of the ventricles into vibration, and the vibrations travel away from the valves. When the vibrations reach the chest wall where the wall is in contact with the heart, sound waves are created that can be heard with the aid of a stethoscope.

      The second heart sound results from vibrations set up in the walls of the pulmonary artery, the aorta, and, to a lesser extent, the ventricles, as the blood reverberates back and forth between the walls of the arteries and the valves after the pulmonary and aortic semilunar valves suddenly close. These vibrations are then heard as a high-frequency sound as the chest wall transforms the vibrations into sound waves. The first heart sound is followed after a short pause by the second. A pause about twice as long comes between the second sound and the beginning of the next cycle. The opening of the valves is silent.

Wall of the heart
      The wall of the heart consists of three distinct layers—the epicardium (outer layer), the myocardium (middle layer), and the endocardium (inner layer). Coronary (coronary circulation) vessels supplying arterial blood to the heart penetrate the epicardium before entering the myocardium. This outer layer, or visceral pericardium, consists of a surface of flattened epithelial (epithelium) (covering) cells resting upon connective tissue.

 The myocardial layer contains the contractile elements of the heart. The bundles of striated muscle fibres present in the myocardium are arranged in a branching pattern and produce a wringing type of movement that efficiently squeezes blood from the heart with each beat. The thickness of the myocardium varies according to the pressure generated to move blood to its destination. The myocardium of the left ventricle, which must drive blood out into the systemic circulation, is, therefore, thickest; the myocardium of the right ventricle, which propels blood to the lungs, is moderately thickened, while the atrial walls are relatively thin.

      The component of the myocardium that causes contraction consists of muscle fibres that are made up of cardiac muscle cells. Each cell contains smaller fibres known as myofibrils that house highly organized contractile units called sarcomeres. The mechanical function arising from sarcomeres is produced by specific contractile proteins (protein) known as actin and myosin (or thin and thick filaments, respectively). The sarcomere, found between two Z lines (or Z discs) in a muscle fibre, contains two populations of actin filaments that project from opposite Z lines in antiparallel fashion and are organized around thick filaments of myosin. As actin slides along crossbridges that project from myosin filaments at regular intervals, each myosin is brought into contact with an adjacent myosin filament. This process shortens the muscle fibre and causes contraction (see muscle).

      Interaction between actin and myosin is regulated by a variety of biological processes that are generally related to the concentration of calcium within the cell. The process of actin sliding over myosin requires large amounts of both calcium and energy. While the contractile machinery occupies about 70 percent of the cardiac cell volume, mitochondria occupy about 25 percent and provide the necessary energy for contraction. To facilitate energy and calcium conductance in cardiac muscle cells, unique junctions called intercalated discs (gap junctions) link the cells together and define their borders. Intercalated discs are the major portal for cardiac cell-to-cell communication, which is required for coordinated muscle contraction and maintenance of circulation.

      Forming the inner surface of the myocardial wall is a thin lining called the endocardium. This layer lines the cavities of the heart, covers the valves and small muscles associated with opening and closing of the valves, and is continuous with the lining membrane of the large blood vessels.

Mark L. Entman Stanley W. Jacob Ed.

Blood supply to the heart
      Because of the watertight lining of the heart (the endocardium) and the thickness of the myocardium, the heart cannot depend on the blood contained in its own chambers for oxygen and nourishment. It possesses a vascular system of its own, called the coronary (coronary artery) arterial system. In the most common distribution, this comprises two major coronary arteries (coronary artery), the right and the left; normally, the left coronary artery divides soon after its origin into two major branches, called the left anterior descending and the circumflex coronary arteries. The right, the left anterior descending, and the left circumflex coronary arteries have many branches and are of almost equal importance. Thus, there are commonly said to be three main functional coronary arteries rather than two.

      The right and left coronary arteries originate from the right and left aortic sinuses (the sinuses of Valsalva), which are bulges at the origin of the ascending aorta immediately beyond, or distal to, the aortic valve. The ostium, or opening, of the right coronary artery is in the right aortic sinus and that of the left coronary artery is in the left aortic sinus, just above the aortic valve ring. There is also a non-coronary sinus of Valsalva, which lies to the left and posteriorly at the origin of the ascending aorta. The left coronary arterial system is more important than the right because it supplies blood to the larger left ventricle, and the dimension of the left coronary ostium is larger than that of the right.

      The right coronary artery has a lumen diameter of about 2.5 millimetres or more. It supplies the right ventricular outflow tract, the sinoatrial node (the principal pacemaker of the heart), the atrioventricular node, and the bulk of the right ventricle, with branches extending into the interventricular septum and joining with arteriolar branches from the left coronary artery more or less where the two ventricles join.

      The main stem of the left coronary artery has a luminal diameter often exceeding 4.5 millimetres and is one of the shortest and most important vessels of the body. Usually, it is between 1 and 2 centimetres in length, but it may have a length of only 2 millimetres before dividing. Sometimes the main left coronary artery may actually be missing, with the left coronary ostium having two separate openings for the left anterior descending and the left circumflex arteries. The main left coronary artery divides into its two branches, the anterior descending and the circumflex, while still in the space between the aorta and pulmonary (pulmonary circulation) artery. The left anterior descending coronary artery usually begins as a continuation of the left main coronary artery, and its size, length, and distribution are key factors in the balance of the supply of blood to the left ventricle and the interventricular septum. There are many branches of the left anterior descending artery; the first and usually the largest septal branch is important because of its prominent role in supplying blood to the septum.

      The left circumflex artery leaves the left main coronary artery to run posteriorly along the atrioventricular groove. It divides soon after its origin into an atrial branch and an obtuse marginal branch. The former branch sometimes has a branch to the sinoatrial node (more usually supplied from the right coronary artery). The obtuse marginal vessel supplies the posterior left ventricular wall in the direction of the apex.

      Venous blood from the heart is carried through veins (vein), which usually accompany the distribution of the distal arteries. These cardiac veins, however, proceed into the atrioventricular grooves anteriorly and posteriorly to form the coronary venous sinus, which opens into the right ventricle immediately below the tricuspid valve.

      In addition to these identifiable anatomic arterial and venous channels, nutritional exchange almost certainly takes place between the endocardial ventricular muscle layers and the blood in the cavity of the ventricles. This is of minor importance and probably is an adaptive system in situations of cardiac muscle pathology.

Michael Francis Oliver

Regulation of heartbeat
      Regular beating of the heart is achieved as a result of the inherent rhythmicity of cardiac muscle (muscle system, human); no nerves are located within the heart itself, and no outside regulatory mechanisms are necessary to stimulate the muscle to contract rhythmically. That these rhythmic contractions originate in the cardiac muscle can be substantiated by observing cardiac development in the embryo (see above (human cardiovascular system)); cardiac pulsations begin before adequate development of nerve fibres. In addition, it can be demonstrated in the laboratory that even fragments of cardiac muscle in tissue culture continue to contract rhythmically. Furthermore, there is no gradation in degree of contraction of the muscle fibres of the heart, as would be expected if they were primarily under nervous control.

      The mere possession of this intrinsic ability is not sufficient, however, to enable the heart to function efficiently. Proper function requires coordination, which is maintained by an elaborate conducting system within the heart that consists primarily of two small, specialized masses of tissue, or nodes, from which impulses originate, and of nervelike conduits for the transmission of impulses, with terminal branches extending to the inner surface of the ventricles.

      Rhythmic cardiac contractions originate with an electrical impulse that travels from the top of the heart in the atria to the bottom of the heart in the ventricles. The impulse is propagated as a wave that travels from cell to cell. Voltage-sensitive protein channels on the surface of the sarcolemma, the membrane that surrounds the muscle fibre, support the flow of current as it relates to the flow of specific ions (ion) (ion-specific channels). These voltage-sensitive channels open and close as a function of the voltage that is sensed on the outer side and inner side (referred to as being “across the membrane,” or transmembrane) of the sarcolemma, between which a difference in electrical potential exists. An electrical potential gradient is created by an excess of negative ions immediately inside the sarcolemma and an equal excess of positive ions on the outside of the sarcolemma (a stage known as the resting potential). When a nerve impulse stimulates ion channels to open, positive ions flow into the cell and cause depolarization, which leads to muscle cell contraction.

      Under resting conditions the heart cell is primarily permeable only to positively charged potassium ions, which slowly leak into the cell. In specialized pacemaking cells, found in the sinoatrial node, the negative resting potential rhythmically drifts toward the positive threshold potential. When the threshold potential is exceeded, depolarization of the cell is triggered, and there is an opening of ion channels that transport sodium and calcium into the cell. This sudden increase in cardiac membrane potential is transmitted from cell to cell, creating a wave of depolarization that functionally represents the excitation signal of the heart. Propagation of the signal rapidly progresses down conduction tissue via specialized atrial cells, the atrioventricular node, and the bundles of His and Purkinje cells and is followed by a slower dispersion of the signal in ventricular muscle cells. The rate of spontaneous depolarization is an important determinant of heart rate.

      Both the excitation and propagation mechanisms are sensitive to alterations in the ion concentration of the extracellular and intracellular fluid, as well as drugs that might alter the carriers or channels associated with these ions. Following the initial depolarization event in cardiac muscle cells, there is a sequence of openings and closures of specific channels that ultimately result in a return to the resting transmembrane potential. This highly orchestrated interaction of different voltage-sensitive channels, and the resultant changes in transmembrane voltage, is termed the cardiac action potential.

      The depolarization event in the cardiac muscle cell also opens a calcium channel, allowing calcium to enter the myocardium. Calcium is an important effector of the coupling between cardiac depolarization (excitation) and cardiac contraction (called “excitation-contraction coupling”). Under normal circumstances, free calcium ion concentration in the cardiac muscle cell is very low. This low concentration is maintained by the presence of an internal membrane system called the sarcoplasmic reticulum that sequesters calcium ions. Upon excitation and depolarization of the cell, the calcium channel opens and admits a small amount of calcium associated with the shift in the membrane potential. This small amount of calcium stimulates the release of additional calcium from calcium-sensitive channels in the sarcoplasmic reticulum, causing the cellular calcium concentration to rise by nearly 100-fold. When the heart is repolarized, the sarcoplasmic reticulum reabsorbs the excess calcium, and the cellular calcium concentration returns to its formerly low level, letting the heart muscle relax.

      Reabsorption of cellular calcium by the sarcoplasmic reticulum is important because it prevents the development of muscle tension. In the resting state, two proteins, troponin and tropomyosin, bind to actin molecules and inhibit interaction between actin and myosin, thereby blocking muscle contraction. When calcium concentration increases during depolarization, it shifts the conformation of troponin and tropomyosin, and actin is able to associate with myosin. As calcium is taken up again by the sarcoplasmic reticulum the myocardial cell relaxes. Factors that control the rise and fall of calcium concentrations in the cardiac muscle cell have profound effects on cardiac function.

 The electrical impulse that is generated by each depolarization of the heart can be characterized and examined with the use of an electrocardiogram. From a clinical standpoint, the electrocardiogram has become useful as a mechanism of diagnosing cardiac disease. The circuitry of the electrocardiogram allows the detection of small changes in voltage that occur rhythmically with cardiac excitation. It was discovered in the early 20th century that these changes could be evaluated by leads (wires) that were placed on the chest, arms, and legs. Potential differences between different sets of leads are examined throughout the cardiac cycle. Ultimately, the readout of the electrocardiogram describes the electrical activation of the heart.

      As a wave of depolarization passes over the atria, the impulse is recorded as the P wave. As it continues through the ventricles, it is registered as the QRS complex. Currents generated as the ventricles recover from the state of depolarization produce the T wave. This repolarization process occurs in the muscle of the ventricles about 0.25 second after depolarization. There are, therefore, both depolarization and repolarization waves represented in the electrocardiogram. The atria repolarize at the same time that the ventricles depolarize; however, the atrial repolarization wave is obscured by the larger QRS wave. The relative timing, size, and direction of these waves are all important diagnostic information in the evaluation of cardiac electrical function and heart disease.

Nervous control of the heart
      Nervous control of the heart is maintained by the parasympathetic fibres in the vagus nerve (parasympathetic) and by the sympathetic nerves. The vagus nerve is the cardiac inhibitor, and the sympathetic nerves are the cardiac excitors. Stimulation of the vagus nerve depresses the rate of impulse formation and atrial contractility and thereby reduces cardiac output and slows the rate of the heart. Parasympathetic stimulation can also produce varying degrees of impaired impulse formation or heart block in diseases of the heart. (In complete heart block the atria and the ventricles beat independently.) Stimulation of the sympathetic nerves increases contractility of both atria and ventricles.

      The cardiac cycle is defined as the time from the end of one heart contraction to the end of the subsequent contraction and consists of a period of relaxation called diastole followed by a period of contraction called systole. During the entire cycle, pressure is maintained in the arteries; however, this pressure varies during the two periods, the normal diastolic pressure being 60 to 80 millimetres of mercury and the normal systolic pressure being 90 to 120 millimetres of mercury.

The blood vessels (blood vessel)
      Because of the need for the early development of a transport system within the embryo, the organs of the vascular system are among the first to appear and to assume their functional role. In fact, this system is established in its basic form by the fourth week of embryonic life. At approximately the 18th day of gestation, cells begin to group together between the outer skin ( ectoderm) and the inner skin ( endoderm) of the embryo. These cells soon become rearranged so that the more peripheral ones join to form a continuous flattened sheet enclosing more centrally placed cells; these cells remain suspended in a fluid medium as primitive blood cells. The tubes then expand and unite to form a network; the primitive blood vessels (blood vessel) thus appear.

      The blood vessels consist of a closed system of tubes that transport blood to all parts of the body and back to the heart. As in any biologic system, structure and function of the vessels are so closely related that one cannot be discussed without the other's being taken into account.

      Arteries (artery) transport blood to body tissues under high pressure, which is exerted by the pumping action of the heart. The heart forces blood into these elastic tubes, which recoil, sending blood on in pulsating waves. It is, therefore, imperative that the vessels possess strong, elastic walls to ensure fast, efficient blood flow to the tissues.

      The wall of an artery consists of three layers. The tunica intima, the innermost layer, consists of an inner surface of smooth endothelium covered by a surface of elastic tissues. The tunica media, or middle coat, is thicker in arteries, particularly in the large arteries, and consists of smooth muscle cells intermingled with elastic fibres. The muscle cells and elastic fibres circle the vessel. In larger vessels the tunica media is composed primarily of elastic fibres. As arteries become smaller, the number of elastic fibres decreases while the number of smooth muscle fibres increases. The tunica adventitia, the outermost layer, is the strongest of the three layers. It is composed of collagenous and elastic fibres. ( collagen is a connective-tissue protein.) The tunica adventitia provides a limiting barrier, protecting the vessel from overexpansion. Also characteristic of this layer is the presence of small blood vessels called the vasa vasorum that supply the walls of larger arteries and veins (vein). In contrast, the inner and middle layers are nourished by diffusion from the blood as it is transported. The thicker, more elastic wall of arteries enables them to expand with the pulse and to regain their original size.

      The transition from artery to arteriole is a gradual one, marked by a progressive thinning of the vessel wall and a decrease in the size of the lumen, or passageway. In arterioles, the tunica intima is still present as a lining covered by a layer of thin longitudinal fibres; however, the tunica media no longer contains elastic fibres and is composed of only a single layer of circular or spiral smooth muscle fibres. The tunica adventitia consists of connective tissue elements.

      The small arteries and arterioles act as control valves through which blood is released into the capillaries (capillary). The strong muscular wall is capable of closing the passageway or permitting it to expand to several times its normal size, thereby vastly altering blood flow to the capillaries. Blood flow is by this device directed to tissues that require it most.

      As the arterioles become smaller in size, the three coats become less and less definite, with the smallest arterioles consisting of little more than endothelium, or lining, surrounded by a layer of smooth muscle. The microscopic capillary tubules consist of a single layer of endothelium that is a continuation of the innermost lining cells of arteries and veins.

      As the capillaries converge, small venules are formed whose function it is to collect blood from the capillary beds (i.e., the networks of capillaries). The venules consist of an endothelial tube supported by a small amount of collagenous tissue and, in the larger venules, by a few smooth muscle fibres as well. As venules continue to increase in size, they begin to exhibit wall structure that is characteristic of arteries, though they are much thinner.

      In veins (vein), which function to conduct blood from the peripheral tissues to the heart, an endothelial lining is surrounded by the tunica media, which contains less muscle and elastic tissue than is found in the arterial wall. The outermost layer, tunica adventitia, is composed chiefly of connective tissue. blood pressure in these vessels is extremely low as compared with that in the arterial system, and blood must exit at an even lower pressure. This creates a need for a special mechanism to keep blood moving on its return to the heart.

      To achieve this, many veins possess a unique system of valves (valve). These valves, formed by semilunar folds in the tunica intima, are present in pairs and serve to direct the flow of blood to the heart, particularly in an upward direction. As blood flows toward the heart, the flaps of the valves flatten against the wall of the vein; they then billow out to block the opening as the pressure of the blood and surrounding tissues fills the valve pocket. These valves are more abundant in the veins of the extremities than in any other parts of the body.

      The veins are more distensible than arteries, and their walls are so constructed as to enable them to expand or contract. A major function of their contractility appears to be to decrease the capacity of the cardiovascular system by constriction of the peripheral vessels in response to the heart's inability to pump sufficient blood.

      Veins tend to follow a course parallel to that of arteries but are present in greater number. Their channels are larger than those of arteries, and their walls are thinner. About 60 percent of the blood volume is in the systemic circulation, and 40 percent is normally present in the veins.

      The pulmonary (pulmonary circulation) circuit consists of the right ventricle; the exiting pulmonary artery and its branches; the arterioles, capillaries, and venules of the lungs (lung); and the pulmonary veins that empty into the left atrium.

      The pulmonary trunk, the common stem of the pulmonary arteries, arises from the upper surface of the right ventricle and extends four to five centimetres beyond this origin before dividing into the right and left pulmonary arteries, which supply the lungs. The pulmonary valve, which has two leaflets, or cusps, guards the opening between the right ventricle and the pulmonary trunk. The trunk is relatively thin-walled for an artery, having walls approximately twice the thickness of the vena cava and one-third that of the aorta. The right and left pulmonary arteries are short but possess a relatively large diameter. The walls are distensible, allowing the vessels to accommodate the stroke volume of the right ventricle, which is a necessary function equal to that of the left ventricle.

      The pulmonary trunk passes diagonally upward to the left across the route of the aorta. Between the fifth and sixth thoracic vertebrae (vertebral column) (at about the level of the bottom of the breastbone), the trunk divides into two branches—the right and left pulmonary arteries—which enter the lungs. After entering the lungs, the branches go through a process of subdivision, the final branches being capillaries. Capillaries surrounding the air sacs (alveoli) of the lungs pick up oxygen and release carbon dioxide. The capillaries carrying oxygenated blood join larger and larger vessels until they reach the pulmonary veins, which carry oxygenated blood from the lungs to the left atrium of the heart.

The arteries
The aorta and its principal branches
      The aorta is the largest vessel in the systemic circuit, arising from the left ventricle. It is commonly said to have three regions: the ascending aorta, the arch of the aorta, and the descending aorta; the latter may be further subdivided into the thoracic and the abdominal aorta.

      Originating from the ascending portion of the aorta are the right and left coronary arteries (coronary artery), which supply the heart with oxygenated blood. Branching from the arch of the aorta are three large arteries named, in order of origin from the heart, the innominate, the left common carotid (carotid artery), and the left subclavian. These three branches supply the head, neck, and arms with oxygenated blood.

      As the innominate (sometimes referred to as the brachiocephalic) artery travels upward toward the clavicle, or collarbone, it divides into the right common carotid and right subclavian arteries. The two common carotid arteries, one branching from the innominate and the other directly from the aorta, then extend in a parallel fashion on either side of the neck to the top of the thyroid (thyroid gland) cartilage (the principal cartilage in the voice box, or larynx), where they divide, each to become an internal and an external carotid artery. The external carotid arteries give off branches that supply much of the head and neck, while the internal carotids are responsible for supplying the forward portion of the brain, the eye and its appendages, and the forehead and nose.

      The two vertebral arteries, one arising as a branch of the innominate and the other as a branch of the left subclavian artery, unite at the base of the brain to form the basilar artery, which in turn divides into the posterior cerebral arteries. The blood supply to the brain is derived mainly from vessels that may be considered as branches of the circle of Willis, which is made up of the two vertebral and the two internal carotid arteries and connecting arteries between them.

      The arms are supplied by the subclavian artery on the left and by the continuation of the innominate on the right. At approximately the border of the first rib, both of these vessels become known as the axillary artery; this, in turn, becomes the brachial artery as it passes down the upper arm. At about the level of the elbow, the brachial artery divides into two terminal branches, the radial and ulnar arteries, the radial passing downward on the distal (thumb) side of the forearm, the ulnar on the medial side. Interconnections (anastomoses) between the two, with branches at the level of the palm, supply the hand and wrist.

      The thoracic (thoracic cavity) (chest) portion of the descending aorta gives off branches that supply the viscera (visceral branches) and the walls surrounding the thoracic cavity (parietal branches). The visceral branches provide blood for the pericardium (human cardiovascular system), lungs, bronchi, lymph nodes (lymph node), and esophagus. The parietal vessels supply the intercostal muscles (the muscles between the ribs (rib)) and the muscles of the thoracic wall; they supply blood to the membrane covering the lungs and lining the thoracic cavity, the spinal cord, the vertebral column, and a portion of the diaphragm.

      As the aorta descends through the diaphragm, it becomes known as the abdominal aorta and again gives off both visceral and parietal branches. Visceral vessels include the celiac, superior mesenteric, and inferior mesenteric, which are unpaired, and the renal and testicular or ovarian, which are paired. The celiac artery arises from the aorta a short distance below the diaphragm and almost immediately divides into the left gastric artery, serving part of the stomach and esophagus; the hepatic artery, which primarily serves the liver; and the splenic artery, which supplies the stomach, pancreas, and spleen.

      The superior mesenteric artery arises from the abdominal aorta just below the celiac artery. Its branches supply the small intestine and part of the large intestine. Arising several centimetres above the termination of the aorta is the inferior mesenteric artery, which branches to supply the lower part of the colon. The renal arteries (renal artery) pass to the kidneys (kidney). The testicular or ovarian arteries supply the testes (testis) in the male and the ovaries (ovary) in the female, respectively.

      Parietal branches of the abdominal aorta include the inferior phrenic, serving the suprarenal (adrenal (adrenal gland)) glands, the lumbar, and the middle sacral arteries. The lumbar arteries are arranged in four pairs and supply the muscles of the abdominal wall, the skin, the lumbar vertebrae, the spinal cord, and the meninges (spinal-cord coverings).

      The abdominal aorta divides into two common iliac arteries, each of which descends laterally and gives rise to external and internal branches. The right and left external iliac arteries are direct continuations of the common iliacs and become known as the femoral arteries after passing through the inguinal region, giving off branches that supply structures of the abdomen and lower extremities.

      At a point just above the knee, the femoral artery continues as the popliteal artery; from this arise the posterior and anterior tibial arteries. The posterior tibial artery is a direct continuation of the popliteal, passing down the lower leg to supply structures of the posterior portion of the leg and foot.

      Arising from the posterior tibial artery a short distance below the knee is the peroneal artery; this gives off branches that nourish the lower leg muscles and the fibula (the smaller of the two bones in the lower leg) and terminate in the foot. The anterior tibial artery passes down the lower leg to the ankle, where it becomes the dorsalis pedis artery, which supplies the foot.

      An impulse can be felt over an artery that lies near the surface of the skin. The impulse results from alternate expansion and contraction of the arterial wall because of the beating of the heart. When the heart pushes blood into the aorta, the blood's impact on the elastic walls creates a pressure wave that continues along the arteries. This impact is the pulse. All arteries have a pulse, but it is most easily felt at points where the vessel approaches the surface of the body.

      The pulse is readily distinguished at the following locations: (1) at the point in the wrist where the radial artery approaches the surface; (2) at the side of the lower jaw where the external maxillary (facial) artery crosses it; (3) at the temple above and to the outer side of the eye, where the temporal artery is near the surface; (4) on the side of the neck, from the carotid artery; (5) on the inner side of the biceps, from the brachial artery; (6) in the groin, from the femoral artery; (7) behind the knee, from the popliteal artery; (8) on the upper side of the foot, from the dorsalis pedis artery.

      The radial artery is most commonly used to check the pulse. Several fingers are placed on the artery close to the wrist joint. More than one fingertip is preferable because of the large, sensitive surface available to feel the pulse wave. While the pulse is being checked, certain data are recorded, including the number and regularity of beats per minute, the force and strength of the beat, and the tension offered by the artery to the finger. Normally, the interval between beats is of equal length.

The veins (vein)
      Venules collect blood from the capillaries and the blood channels known as sinusoids (sinusoid) and unite to form progressively larger veins that terminate as the great veins, or venae cavae. In the extremities there are superficial and deep veins; the superficial lie just under the skin and drain the skin and superficial fasciae (sheets of fibrous tissue), while the deep veins accompany the principal arteries of the extremities and are similarly named. Interconnections between the superficial and the deep veins are frequent.

      Venous blood enters the right atrium from three sources: the heart muscle by way of the coronary sinus; the upper body by way of the superior vena cava (vena cava); and the lower body by way of the inferior vena cava (vena cava).

Superior vena cava and its tributaries
      Tributaries from the head and neck, the arms, and part of the chest unite to form the superior vena cava. Venous channels called venous sinuses (venous sinus) lie between the two layers of the dura mater, the outer covering of the brain; they possess no valves. Venous drainage of the brain is effected by these sinuses and communicating vessels. The internal jugular vein is a continuation of this system downward through the neck; it receives blood from parts of the face, neck, and brain. At approximately the level of the collarbone, each unites with the subclavian vein of that side to form the innominate veins.

      The external jugular vein is formed by the union of its tributaries near the angle of the lower jaw, or mandible. It drains some of the structures of the head and neck and pours its contents along with the subclavian into the innominate vein of the same side. All of the veins of the arm are tributaries of the subclavian vein of that side. They are found in both superficial and deep locations and possess valves. Most of the deep veins are arranged in pairs with cross connections between them.

      Venous drainage of the hand is accomplished superficially by small anastomosing (interconnecting) veins that unite to form the cephalic vein, coursing up the radial (thumb) side of the forearm, and the basilic vein, running up the ulnar side of the forearm and receiving blood from the hand, forearm, and arm. The deep veins of the forearm include the radial veins, continuations of deep anastomosing veins of the hand and wrist, and the ulnar veins, both veins following the course of the associated artery. The radial and ulnar veins converge at the elbow to form the brachial vein; this, in turn, unites with the basilic vein at the level of the shoulder to produce the axillary vein. At the outer border of the first rib, the axillary vein becomes the subclavian vein, the terminal point of the venous system characteristic of the upper extremity.

      The subclavian, external jugular, and internal jugular veins all converge to form the innominate vein. The right and left innominate veins terminate in the superior vena cava, which opens into the upper posterior portion of the right atrium.

      In addition to the innominate veins, the superior vena cava receives blood from the azygous vein and small veins from the mediastinum (the region between the two lungs) and the pericardium. Most of the blood from the back and from the walls of the chest and abdomen drains into veins lying alongside the vertebral bodies (the weight-bearing portions of the vertebrae). These veins form what is termed the azygous system, which serves as a connecting link between the superior and inferior vena cava. The terminal veins of this system are the azygous, hemiazygous, and accessory hemiazygous veins. At the level of the diaphragm, the right ascending lumbar vein continues upward as the azygous vein, principal tributaries of which are the right intercostal veins, which drain the muscles of the intercostal spaces. It also receives tributaries from the esophagus, lymph nodes, pericardium, and right lung, and it enters into the superior vena cava at about the level of the fourth thoracic vertebra.

      The left side of the azygous system varies greatly among individuals. Usually the hemiazygous vein arises just below the diaphragm as a continuation of the left ascending lumbar vein and terminates in the azygous vein. Tributaries of the hemiazygous drain the intercostal muscles, the esophagus, and a portion of the mediastinum. The accessory hemiazygous usually extends downward as a continuation of the vein of the fourth intercostal space, receiving tributaries from the left intercostal spaces and the left bronchus. It empties into the azygous vein slightly above the entrance of the hemiazygous.

Inferior vena cava and its tributaries
      The inferior vena cava is a large, valveless, venous trunk that receives blood from the legs, the back, and the walls and contents of the abdomen and pelvis.

      The foot is drained primarily by the dorsal venous arch, which crosses the top of the foot not far from the base of the toes. The arch is connected with veins that drain the sole. Superficially the lower leg is drained by the large and small saphenous veins, which are continuations of the dorsal venous arch. The small saphenous vein extends up the back of the lower leg to terminate usually in the popliteal vein. There is some interconnection with deep veins and with the great saphenous vein. The latter vein, the longest in the body, extends from the dorsal venous arch up the inside of the lower leg and thigh, receiving venous branches from the knee and thigh area and terminating in the femoral vein.

      Most blood from the lower extremity returns by way of the deep veins. These include the femoral and popliteal veins and the veins accompanying the anterior and posterior tibial and peroneal arteries. The anterior and posterior tibial veins originate in the foot and join at the level of the knee to form the popliteal vein; the latter becomes the femoral vein as it continues its extension through the thigh.

      At the level of the inguinal ligament (which is at the anterior, diagonal border between the trunk and the thigh), the femoral vein becomes known as the external iliac vein; the latter unites with the internal iliac vein to form the common iliac vein. The internal iliac vein drains the pelvic walls, viscera, external genitalia, buttocks, and a portion of the thigh. Through the paired common iliac veins, the legs and most of the pelvis are drained. The two common iliacs then unite at a level above the coccyx (the lowest bone in the spine) to become the inferior vena cava. As it courses upward through the abdomen, the inferior vena cava receives blood from the common iliacs and from the lumbar, renal, suprarenal, and hepatic veins before emptying into the right atrium.

      The pairs of lumbar veins (which drain blood from the loins and abdominal walls) are united on each side by a vertical connecting vein, the ascending lumbar vein; the right ascending lumbar vein continues as the azygous and the left as the hemiazygous. These veins usually enter separately into the inferior vena cava.

      Renal veins lie in front of the corresponding renal artery; the right renal vein receives tributaries exclusively from the kidney, while the left receives blood from a number of other organs as well. The right suprarenal vein terminates directly in the inferior vena cava as does the right phrenic, above the gonadal vein. Two or three short hepatic trunks empty into the inferior vena cava as it passes through the diaphragm.

      The portal system may be described as a specialized portion of the systemic circulatory system (systemic circulation). Although it originates in capillaries (capillary), the portal system is unique from other vessels in that it also terminates in a capillary-like vascular bed, located in the liver. The blood from the spleen, stomach, pancreas, and intestine first passes through the liver before it moves on to the heart. Blood flowing to the liver comes from the hepatic artery (20 percent) and the portal vein (80 percent); blood leaving the liver flows through the hepatic vein and then empties into the inferior vena cava. The hepatic arterial blood supplies oxygen requirements for the liver. Blood from the abdominal viscera, particularly the intestinal tract, passes into the portal vein and then into the liver. Substances in the portal blood are processed by the liver (see digestive system, human).

Venous pulmonary system
      From the pulmonary capillaries, in which blood takes on oxygen and gives up carbon dioxide, the oxygenated blood in veins is collected first into venules and then into progressively larger veins; it finally flows through four pulmonary veins, two from the hilum of each lung. (The hilum is the point of entry on each lung for the bronchus, blood vessels, and nerves.) These veins then pass to the left atrium, where their contents are poured into the heart.

The capillaries (capillary)
      The vast network of some 10,000,000,000 microscopic capillaries functions to provide a method whereby fluids, nutrients, and wastes are exchanged between the blood and the tissues. Even though microscopic in size, the largest capillary being approximately 0.2 millimetre in diameter (about the width of the tip of a pin), the great network of capillaries serves as a reservoir normally containing about one-sixth of the total circulating blood volume. The number of capillaries in active tissue, such as muscle, liver, kidney, and lungs, is greater than the number in tendon or ligament; in addition, the cornea of the eye, epidermis, and hyaline cartilage (semitransparent cartilage such as is found in joints) are devoid of capillaries.

      The interconnecting network of capillaries into which the arterioles empty is characterized not only by microscopic size but also by extremely thin walls only one cell thick. The vessels are simply tubular continuations of the inner lining cells of the larger vessels, normally uniform in size, usually three to four endothelial cells in circumference, except toward the venous terminations, where they become slightly wider, four to six cells in circumference. A thin membrane, called a basement membrane, surrounds these cells and serves to maintain the integrity of the vessel.

      A single capillary unit consists of a branching and interconnecting (anastomosing) network of vessels, each averaging 0.5 to 1 millimetre in length. The wall of the capillary is extremely thin and acts as a semipermeable membrane that allows substances containing small molecules, such as oxygen, carbon dioxide, water, fatty acids (fatty acid), glucose, and ketones (ketone), to pass through the membrane. Oxygen and nutritive material pass into the tissues through the wall at the arteriolar end of the capillary unit; carbon dioxide and waste products move through the membrane into the vessel at the venous end of the capillary bed. Constriction and dilation of the arterioles is primarily responsible for regulating the flow of blood into the capillaries. Muscular gatekeepers, or sphincters (sphincter muscle), in the capillary unit itself, however, serve to direct the flow to those areas in greatest need.

      There are three modes of transport across the cellular membrane of the capillary wall. Substances soluble in the lipid (fatty) membrane of the capillary cells can pass directly through these membranes by a process of diffusion. Some substances needed by the tissues and soluble in water but completely insoluble in the lipid membrane pass through minute water-filled passageways, or pores, in the membranes by a process called ultrafiltration. Only 1/1,000th of the surface area of capillaries is represented by these pores. Other substances, such as cholesterol, are transported by specific receptors in the endothelium.

Human fetal circulation
      In the fetus, oxygenated blood is carried from the placenta to the fetus by the umbilical vein. It then passes to the inferior vena cava of the fetus by way of a vessel called the ductus venosus. From the inferior vena cava, the blood enters the right atrium, then passes through the foramen ovale into the left atrium; from there it moves into the left ventricle and out through the aorta, which pumps the oxygenated blood to the head and upper extremities. Blood from the upper extremities returns via the superior vena cava into the right atrium, where it is largely deflected into the right ventricle.

      From the right ventricle, a portion of the blood flows into the pulmonary artery to the lungs. The largest fraction flows through an opening, the ductus arteriosus, into the aorta. It enters the aorta beyond the point at which the blood of the head leaves. Some of the blood supplies the lower portion of the body. The remainder returns to the placenta via the umbilical arteries, which branch off from the internal iliac arteries.

      The changes that take place at birth and that permit routing of the blood through the pulmonary system (pulmonary circulation) instead of the umbilical vessels have been described above (human cardiovascular system) in the section on the origin and development of the heart.

Evaluating the cardiovascular system
      Certain diagnostic (diagnosis) techniques with respect to the heart and blood vessels (blood vessel) are important factors in determining the degree of disease and their appropriate medical and surgical treatment.

Invasive techniques
Right-heart catheterization
 Right-heart catheterization (cardiac catheterization) is performed by insertion of a catheter (a long tube) into the cubital vein (at the bend of the elbow), the saphenous vein (in the inner thigh), or the femoral vein (at the groin). The catheter, which is opaque to X-ray, is advanced into the right atrium, right ventricle, and pulmonary artery under fluoroscopy. This procedure makes it possible to measure pressure and oxygen saturation in the right heart chamber itself and thus to diagnose abnormalities in the valves.

Left-heart catheterization
      Left-heart catheterization is accomplished by introducing a catheter into the brachial or femoral artery (in the upper arm and thigh, respectively) and advancing it through the aorta across the aortic valve and into the left ventricle. Mitral and aortic valvular defects and myocardial disease can be evaluated by this technique.

angiocardiography and arteriography
  angiocardiography permits direct visualization of the chambers and great vessels of the heart from injections of dyes that are opaque to X-rays. Anatomic defects, such as congenital and acquired lesions, can be detected readily. Left ventriculography (X-ray pictures of the left ventricle) provides information about the synchrony and adequacy of the forces of contraction in areas of the left ventricle. Arteriography (angiography) (X-ray pictures of an artery after the injection of dyes that are opaque to X-rays) of the coronary arteries (coronary artery) permits identification, localization, and assessment of the extent of obstructive lesions within these arteries. It is the most important means of defining the presence and severity of coronary atherosclerosis and, in conjunction with left ventriculography, the related state of myocardial function. Although invasive techniques involving left ventricular catheterization and radio contrast angiocardiography and arteriography provide reliable measurements of ejection fraction and regional formation, they have limited applications.

Noninvasive techniques
      The term echocardiography refers to a group of tests that use ultrasound (sound waves above frequencies audible to humans) to examine the heart and record information in the form of echoes, or reflected sonic waves. M-mode echocardiography records the amplitude and the rate of motion of moving objects, such as valves, along a single line with great accuracy. M-mode echocardiography, however, does not permit effective evaluation of the shape of cardiac structures, nor does it depict lateral motion (i.e., motion perpendicular to the ultrasonic beam). Real-time (cross-sectional or two-dimensional) echocardiography depicts cardiac shape and lateral movement not available in M-mode echocardiography by moving the ultrasonic beam very rapidly, and such recording may be displayed on film or videotape. New techniques allow measurement by ultrasonography of rates of flow and pressures, for example, across heart valves.

      Radionuclide imaging (radioactive nuclides) provides a safe, quantitative evaluation of cardiac function and a direct measurement of myocardial blood flow and myocardial metabolism. Radionuclide imaging is used to evaluate the temporal progress of cardiac disease, hemodynamics, and the extent of myocardial damage during and after infarction and to detect pulmonary infarction following emboli (embolism). The primary requirement of radionuclide imaging is that the bolus of radionuclide should remain within the blood vessels during its first passage through the right and left sides of the heart. The second requirement is that the physical properties of the radionuclide be satisfactory with respect to the instrumentation being used.

      The radionuclide used in virtually all phases of radionuclide imaging is technetium-99 (technetium). It has the disadvantage of a long half-life (six hours), however, and other radionuclides with shorter half-lives are also used. These radionuclides all emit gamma rays (gamma ray), and a scintillation camera is used to detect gamma-ray emission. The data are assessed with the R wave of the electrocardiogram (human cardiovascular system) as a time marker for the cardiac cycle. Radionuclide cineangiography is a further development of radionuclide imaging. These techniques are used to assess myocardial damage, left ventricular function, valve regurgitation, and, with the use of radionuclide potassium analogues, myocardial perfusion.

      There are techniques that measure metabolism in the myocardium using the radiotracer method (i.e., a radioactive isotope replaces a stable element in a compound, which is then followed as it is distributed through the body). positron emission tomography uses positron radionuclides that can be incorporated into true metabolic substrates and consequently can be used to chart the course of selected metabolic pathways, such as myocardial glucose uptake and fatty-acid (fatty acid) metabolism. Magnetic resonance imaging (MRI; also called nuclear magnetic resonance [NMR]), also allows high resolution tomographic (one-plane) and three-dimensional imaging of tissues. Magnetic resonance imaging uses magnetic fields and radio frequencies to penetrate bone and obtain clear images of the underlying tissues.

Michael Francis Oliver Mark L. Entman Stanley W. Jacob

Additional Reading
General accounts and elementary descriptions of circulatory systems are found in many biology textbooks, including the following: Raymond F. Oram, Biology: Living Systems, 5th ed. (1986); Karen Arms and Pamela S. Camp, Biology, 3rd ed. (1986); and Paul B. Weisz and Richard N. Keogh, The Science of Biology, 5th ed. (1982). Textbooks dealing with animal structure at a more advanced level include the following: Ralph M. Buchsbaum, Animals Without Backbones, 3rd ed. (1987); Robert D. Barnes, Invertebrate Zoology, 5th ed. (1987); Alfred Sherwood Romer and Thomas S. Parsons, The Vertebrate Body, 6th ed. (1986); and Charles K. Weichert, Anatomy of the Chordates, 4th ed. (1970); Knut Schmidt-Nielsen, Animal Physiology: Adaptation and Environment, 3rd ed. (1983); and Milton Hildebrand, Analysis of Vertebrate Structure, 2nd ed. (1982).For the history of circulation studies, see Helen Rapson, The Circulation of Blood (1982); David J. Furley and J.S. Wilkie (eds.), Galen on Respiration and the Arteries (1984); The Selected Writings of William Gilbert, Galileo Galilei, William Harvey (1952), in “The Great Books of the Western World” series; Fredrick A. Willius and Thomas J. Dry, A History of the Heart and the Circulation (1948); and Alfred P. Fishman and Dickinson W. Richards, Circulation of the Blood: Men and Ideas (1964, reprinted 1982). Special studies of circulation include Donald A. McDonald, Blood Flow in Arteries, 2nd ed. (1974); David I. Abramson and Philip B. Dobrin (eds.), Blood Vessels and Lymphatics in Organ Systems (1984); Colin L. Schwartz, Nicholas T. Werthessen, and Stewart Wolf, Structure and Function of the Circulation, 3 vol. (1980–81); and Jerry Franklin Green, Fundamental Cardiovascular and Pulmonary Physiology, 2nd ed. (1987). Stanley W. Jacob, Clarice Ashworth Francone, and Walter J. Lossow, Structure and Function in Man, 5th ed. (1982); and Gary A. Thibodeau, Anatomy and Physiology (1987), are basic texts. Arthur C. Guyton, Human Physiology and Mechanisms of Disease, 4th ed. (1987), is a technical description of the physiology of cardiac muscle, heart function, and hemodynamics. Also see Peter F. Cohn, Clinical Cardiovascular Physiology (1985); James J. Smith and John P. Kampine, Circulatory Physiology: The Essentials (1984); Harvey V. Sparks, Jr., and Thom W. Rooke, Essentials of Cardiovascular Physiology (1987); and Peter Harris and Donald Heath, The Human Pulmonary Circulation, 3rd ed. (1986).Michael Francis Oliver

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