- Compare and contrast the organization/function of circulatory systems, including gastrovascular cavity, open, closed, single, and double systems
- Identify and describe the functions of different types of blood vessels (artery, arteriole, capillary, venule, vein), including their basic structure
- Describe and identify the functions of the different components of blood
- Describe the process of gas, nutrient, and fluid exchange between capillaries and tissues
Types of Circulatory Systems
The information below was adapted from OpenStax Biology 40.1
The circulatory system is the primary method used to transport nutrients and gases through the body. Simple diffusion allows some water, nutrient, waste, and gas exchange in animals that are only a few cell layers thick; however, bulk flow is the only method by which the entire body of larger, more complex organisms is accessed.
Circulatory System Architecture
The circulatory system is effectively a network of cylindrical vessels: the arteries, veins, and capillaries that emanate from a pump, the heart. In all vertebrate organisms, as well as some invertebrates, this is a closed-loop system, in which the blood is not free in a cavity. In a closed circulatory system, blood is contained inside blood vessels and circulates unidirectionally from the heart around the systemic circulatory route, then returns to the heart again.
As opposed to a closed system, arthropods– including insects, crustaceans, and most mollusks– have an ‘open’ circulatory system. In an open circulatory system, the blood is not enclosed in blood vessels but is pumped into an open cavity called a hemocoel and is called hemolymph because the blood mixes with the interstitial fluid. As the heart beats and the animal moves, the hemolymph circulates around the organs within the body cavity and then reenters the hearts through openings called ostia. This movement allows for nutrient exchange, and in some organisms lacking direct gas exchange sites, a basic mechanism to transport gasses beyond the exchange site. Because the gas exchange in many open-circulatory systems tends to be relatively low for metabolically-active organs and tissues, a tradeoff exists between this system and the much more energy-consuming, harder-to-maintain closed system.
In (a) closed circulatory systems, the heart pumps blood through vessels that are separate from the interstitial fluid of the body. Most vertebrates and some invertebrates, like this annelid earthworm, have a closed circulatory system. In (b) open circulatory systems, a fluid called hemolymph is pumped through a blood vessel that empties into the body cavity. Hemolymph returns to the blood vessel through openings called ostia. Arthropods like this bee and most mollusks have open circulatory systems.
Circulatory System Variation in Animals
The circulatory system varies from simple systems in invertebrates to more complex systems in vertebrates. The simplest animals, such as the sponges (Porifera) and rotifers (Rotifera), do not need a circulatory system because diffusion allows adequate exchange of water, nutrients, and waste, as well as dissolved gases. Organisms that are more complex but still only have two layers of cells in their body plan, such as jellies (Cnidaria) and comb jellies (Ctenophora) also use diffusion through their epidermis and internally through the gastrovascular compartment. Both their internal and external tissues are bathed in an aqueous environment and exchange fluids by diffusion on both sides. Exchange of fluids is assisted by the pulsing of the jellyfish body.
Simple animals consisting of a single cell layer such as the (a) sponge or only a few cell layers such as the (b) jellyfish do not have a circulatory system. Instead, gases, nutrients, and wastes are exchanged by diffusion.
For more complex organisms, diffusion is not efficient for cycling gases, nutrients, and waste effectively through the body; therefore, more complex circulatory systems evolved. In an open system, an elongated beating heart pushes the hemolymph through the body and muscle contractions help to move fluids. The larger more complex crustaceans, including lobsters, have developed arterial-like vessels to push blood through their bodies, and the most active mollusks, such as squids, have evolved a closed circulatory system and are able to move rapidly to catch prey. Closed circulatory systems are a characteristic of vertebrates; however, there are significant differences in the structure of the heart and the circulation of blood between the different vertebrate groups due to adaptation during evolution and associated differences in anatomy. The figure below illustrates the basic circulatory systems of some vertebrates: fish, amphibians, reptiles, and mammals.
(a) Fish have the simplest circulatory systems of the vertebrates: blood flows unidirectionally from the two-chambered heart through the gills and then the rest of the body. (b) Amphibians have two circulatory routes: one for oxygenation of the blood through the lungs and skin, and the other to take oxygen to the rest of the body. The blood is pumped from a three-chambered heart with two atria and a single ventricle. (c) Reptiles also have two circulatory routes; however, blood is only oxygenated through the lungs. The heart is three chambered, but the ventricles are partially separated so some mixing of oxygenated and deoxygenated blood occurs except in crocodilians and birds. (d) Mammals and birds have the most efficient heart with four chambers that completely separate the oxygenated and deoxygenated blood; it pumps only oxygenated blood through the body and deoxygenated blood to the lungs.
Fish have a single circuit for blood flow and a two-chambered heart that has only a single atrium and a single ventricle. The atrium collects blood that has returned from the body and the ventricle pumps the blood to the gills where gas exchange occurs and the blood is re-oxygenated; this is called gill circulation. The blood then continues through the rest of the body before arriving back at the atrium; this is called systemic circulation. This unidirectional flow of blood produces a gradient of oxygenated to deoxygenated blood around the fishâ€™s systemic circuit. The result is a limit in the amount of oxygen that can reach some of the organs and tissues of the body, reducing the overall metabolic capacity of fish.
In amphibians, reptiles, birds, and mammals, blood flow is directed in two circuits: one through the lungs and back to the heart, which is called pulmonary circulation, and the other throughout the rest of the body and its organs including the brain (systemic circulation). In amphibians, gas exchange also occurs through the skin during pulmonary circulation and is referred to as pulmocutaneous circulation.
Amphibians have a three-chambered heart that has two atria and one ventricle rather than the two-chambered heart of fish. The two atria (superior heart chambers) receive blood from the two different circuits (the lungs and the systems), and then there is some mixing of the blood in the heart’s ventricle (inferior heart chamber), which reduces the efficiency of oxygenation. The advantage to this arrangement is that high pressure in the vessels pushes blood to the lungs and body. The mixing is mitigated by a ridge within the ventricle that diverts oxygen-rich blood through the systemic circulatory system and deoxygenated blood to the pulmocutaneous circuit. For this reason, amphibians are often described as having double circulation.
Most reptiles also have a three-chambered heart similar to the amphibian heart that directs blood to the pulmonary and systemic circuits. However, the ventricle is divided more effectively by a partial septum, which results in less mixing of oxygenated and deoxygenated blood. Some reptiles (alligators and crocodiles) are the most “primitive” animals to exhibit a four-chambered heart. Crocodilians have a unique circulatory mechanism where the heart shunts blood from the lungs toward the stomach and other organs during long periods of submergence, for instance, while the animal waits for prey or stays underwater waiting for prey to rot. One adaptation includes two main arteries that leave the same part of the heart: one takes blood to the lungs and the other provides an alternate route to the stomach and other parts of the body. Two other adaptations include a hole in the heart between the two ventricles, called the foramen of Panizza, which allows blood to move from one side of the heart to the other, and specialized connective tissue that slows the blood flow to the lungs.
In mammals and birds, the heart is divided completely into four chambers: two atria and two ventricles. Oxygenated blood is fully separated from deoxygenated blood, which improves the efficiency of double circulation and is probably required for supporting the warm-blooded lifestyle of mammals and birds. The four-chambered heart of birds and mammals evolved independently from a three-chambered heart. The independent evolution of the same or a similar biological trait is referred to as convergent evolution.
This video gives an overview of the different types of circulatory systems in different types of animals:
Function and Composition of Blood
The information below was adapted from OpenStax Biology 40.2
Hemoglobin is responsible for distributing oxygen, and to a lesser extent, carbon dioxide, throughout the circulatory systems of humans, vertebrates, and many invertebrates. The blood is more than the proteins, though. Blood is actually a term used to describe the liquid that moves through the vessels and includes plasma (the liquid portion, which contains water, proteins, salts, lipids, and glucose) and the cells (red and white cells) and cell fragments called platelets. Blood plasma is actually the dominant component of blood and contains the water, proteins, electrolytes, lipids, and glucose. The cells are responsible for carrying the gases (red cells) and immune response (white). The platelets are responsible for blood clotting. Interstitial fluid that surrounds cells is separate from the blood, but in hemolymph, they are combined. In humans, cellular components make up approximately 45 percent of the blood and the liquid plasma 55 percent. Blood is 20 percent of a person’s extracellular fluid and eight percent of weight.
The Role of Blood in the Body
Blood, like the human blood illustrated below, is important for regulation of the body’s systems and homeostasis. Blood helps maintain homeostasis by stabilizing pH, temperature, osmotic pressure, and by eliminating excess heat. Blood supports growth by distributing nutrients and hormones, and by removing waste. Blood plays a protective role by transporting clotting factors and platelets to prevent blood loss and transporting the disease-fighting agents or white blood cells to sites of infection.
The cells and cellular components of human blood are shown. Red blood cells deliver oxygen to the cells and remove carbon dioxide. White blood cellsâ€”including neutrophils, monocytes, lymphocytes, eosinophils, and basophilsâ€”are involved in the immune response. Platelets form clots that prevent blood loss after injury.
Red Blood Cells
Red blood cells, or erythrocytes (erythro- = “red”; -cyte = “cell”), are specialized cells that circulate through the body delivering oxygen to cells; they are formed from stem cells in the bone marrow. In mammals, red blood cells are small biconcave cells that at maturity do not contain a nucleus or mitochondria and are only 7â€“8 Âµm in size. In birds and non-avian reptiles, a nucleus is still maintained in red blood cells.
The red coloring of blood comes from the iron-containing protein hemoglobin. The principal job of this protein is to carry oxygen, but it also transports carbon dioxide as well. Hemoglobin is packed into red blood cells at a rate of about 250 million molecules of hemoglobin per cell. Each hemoglobin molecule binds four oxygen molecules so that each red blood cell carries one billion molecules of oxygen. There are approximately 25 trillion red blood cells in the five liters of blood in the human body, which could carry up to 25 sextillion (25 * 1021) molecules of oxygen in the body at any time. In mammals, the lack of organelles in erythrocytes leaves more room for the hemoglobin molecules, and the lack of mitochondria also prevents use of the oxygen for metabolic respiration. Only mammals have anucleated red blood cells, and some mammals (camels, for instance) even have nucleated red blood cells. The advantage of nucleated red blood cells is that these cells can undergo mitosis. Anucleated red blood cells metabolize anaerobically (without oxygen), making use of a primitive metabolic pathway to produce ATP and increase the efficiency of oxygen transport.
Not all organisms use hemoglobin as the method of oxygen transport. Invertebrates that utilize hemolymph rather than blood use different pigments to bind to the oxygen. These pigments use copper or iron to the oxygen. Invertebrates have a variety of other respiratory pigments. Hemocyanin, a blue-green, copper-containing protein is found in mollusks, crustaceans, and some of the arthropods. Chlorocruorin, a green-colored, iron-containing pigment is found in four families of polychaete tubeworms. Hemerythrin, a red, iron-containing protein is found in some polychaete worms and annelids. Despite the name, hemerythrin does not contain a heme group and its oxygen-carrying capacity is poor compared to hemoglobin.
In most vertebrates, (a) hemoglobin delivers oxygen to the body and removes some carbon dioxide. Hemoglobin is composed of four protein subunits, two alpha chains and two beta chains, and a heme group that has iron associated with it. The iron reversibly associates with oxygen, and in so doing is oxidized from Fe2+ to Fe3+. In most mollusks and some arthropods, (b) hemocyanin delivers oxygen. Unlike hemoglobin, hemolymph is not carried in blood cells, but floats free in the hemolymph. Copper instead of iron binds the oxygen, giving the hemolymph a blue-green color. In annelids, such as the earthworm, and some other invertebrates, (c) hemerythrin carries oxygen. Like hemoglobin, hemerythrin is carried in blood cells and has iron associated with it, but despite its name, hemerythrin does not contain heme.
The small size and large surface area of red blood cells allows for rapid diffusion of oxygen and carbon dioxide across the plasma membrane. In the lungs, carbon dioxide is released and oxygen is taken in by the blood. In the tissues, oxygen is released from the blood and carbon dioxide is bound for transport back to the lungs. Studies have found that hemoglobin also binds nitrous oxide (NO). NO is a vasodilator that relaxes the blood vessels and capillaries and may help with gas exchange and the passage of red blood cells through narrow vessels. Nitroglycerin, a heart medication for angina and heart attacks, is converted to NO to help relax the blood vessels and increase oxygen flow through the body.
A characteristic of red blood cells is their glycolipid and glycoprotein coating; these are lipids and proteins that have carbohydrate molecules attached. In humans, the surface glycoproteins and glycolipids on red blood cells vary between individuals, producing the different blood types, such as A, B, and O. Red blood cells have an average lifespan of 120 days, at which time they are broken down and recycled in the liver and spleen by phagocytic macrophages, a type of white blood cell.
White Blood Cells
White blood cells, also called leukocytes (leuko = white), make up approximately one percent by volume of the cells in blood. The role of white blood cells is very different than that of red blood cells: they are primarily involved in the immune response to identify and target pathogens, such as invading bacteria, viruses, and other foreign organisms. White blood cells are formed continually; some only live for hours or days, but some live for years.
The morphology of white blood cells differs significantly from red blood cells. They have nuclei and do not contain hemoglobin. The different types of white blood cells are identified by their microscopic appearance after histologic staining, and each has a different specialized function. The two main groups are the granulocytes, which include the neutrophils, eosinophils, and basophils, and the agranulocytes, which include the monocytes and lymphocytes.
(a) Granulocytesâ€”including neutrophils, eosinophils and basophilsâ€”are characterized by a lobed nucleus and granular inclusions in the cytoplasm. Granulocytes are typically first-responders during injury or infection. (b) Agranulocytes include lymphocytes and monocytes. Lymphocytes, including B and T cells, are responsible for adaptive immune response. Monocytes differentiate into macrophages and dendritic cells, which in turn respond to infection or injury.
Platelets and Coagulation Factors
Blood must clot to heal wounds and prevent excess blood loss. Small cell fragments called platelets (thrombocytes) are attracted to the wound site where they adhere by extending many projections and releasing their contents. These contents activate other platelets and also interact with other coagulation factors, which convert fibrinogen, a water-soluble protein present in blood serum into fibrin (a non-water soluble protein), causing the blood to clot. Many of the clotting factors require vitamin K to work, and vitamin K deficiency can lead to problems with blood clotting. Many platelets converge and stick together at the wound site forming a platelet plug (also called a fibrin clot). The plug or clot lasts for a number of days and stops the loss of blood. Platelets are formed from the disintegration of larger cells called megakaryocytes. For each megakaryocyte, 2000-3000 platelets are formed with 150,000 to 400,000 platelets present in each cubic millimeter of blood. Each platelet is disc shaped and 2-4 ¼m in diameter. They contain many small vesicles but do not contain a nucleus.
(a) Platelets are formed from large cells called megakaryocytes. The megakaryocyte breaks up into thousands of fragments that become platelets. (b) Platelets are required for clotting of the blood. The platelets collect at a wound site in conjunction with other clotting factors, such as fibrinogen, to form a fibrin clot that prevents blood loss and allows the wound to heal.
Functions and Types of Blood Vessels
The blood from the heart is carried through the body by a complex network of blood vessels. Arteries take blood away from the heart. The main artery is the aorta that branches into major arteries that take blood to different limbs and organs. These major arteries include the carotid artery that takes blood to the brain, the brachial arteries that take blood to the arms, and the thoracic artery that takes blood to the thorax and then into the hepatic, renal, and gastric arteries for the liver, kidney, and stomach, respectively. The iliac artery takes blood to the lower limbs. The major arteries diverge into minor arteries, and then smaller vessels called arterioles, to reach more deeply into the muscles and organs of the body.
The major human arteries and veins are shown. (credit: modification of work by Mariana Ruiz Villareal)
Arterioles diverge into capillary beds. Capillary beds contain a large number (10 to 100) of capillaries that branch among the cells and tissues of the body. Capillaries are narrow-diameter tubes that can fit red blood cells through in single file and are the sites for the exchange of nutrients, waste, and oxygen with tissues at the cellular level. Fluid also crosses into the interstitial space from the capillaries. The capillaries converge again into venules that connect to minor veins that finally connect to major veins that take blood high in carbon dioxide back to the heart. Veins are blood vessels that bring blood back to the heart. The major veins drain blood from the same organs and limbs that the major arteries supply. Fluid is also brought back to the heart via the lymphatic system.
The structure of the different types of blood vessels reflects their function or layers. There are three distinct layers, or tunics, that form the walls of blood vessels. The first tunic is a smooth, inner lining of endothelial cells that are in contact with the red blood cells. The endothelial tunic is continuous with the endocardium of the heart. In capillaries, this single layer of cells is the location of diffusion of oxygen and carbon dioxide between the endothelial cells and red blood cells, as well as the exchange site via endocytosis and exocytosis. The movement of materials at the site of capillaries is regulated by vasoconstriction, narrowing of the blood vessels, and vasodilation, widening of the blood vessels; this is important in the overall regulation of blood pressure.
Veins and arteries both have two further tunics that surround the endothelium: the middle tunic is composed of smooth muscle and the outermost layer is connective tissue (collagen and elastic fibers). The elastic connective tissue stretches and supports the blood vessels, and the smooth muscle layer helps regulate blood flow by altering vascular resistance through vasoconstriction and vasodilation. The arteries have thicker smooth muscle and connective tissue than the veins to accommodate the higher pressure and speed of freshly pumped blood. The veins are thinner walled as the pressure and rate of flow are much lower. In addition, veins are structurally different than arteries in that veins have valves to prevent the backflow of blood. Because veins have to work against gravity to get blood back to the heart, contraction of skeletal muscle assists with the flow of blood back to the heart.
Arteries and veins consist of three layers: an outer tunica externa, a middle tunica media, and an inner tunica intima. Capillaries consist of a single layer of epithelial cells, the tunica intima. (credit: modification of work by NCI, NIH)
This video describes the structure and function of different types of blood vessels:
Gas, Nutrient, and Fluid Exchange Across Blood Vessels
The information below was adapted from OpenStax Biology 40.4
Blood is pushed through the body by the action of the pumping heart. With each rhythmic pump, blood is pushed under high pressure and velocity away from the heart, initially along the main artery, the aorta. In the aorta, the blood travels at 30 cm/sec. As blood moves into the arteries, arterioles, and ultimately to the capillary beds, the rate of movement slows dramatically to about 0.026 cm/sec, one-thousand times slower than the rate of movement in the aorta. While the diameter of each individual arteriole and capillary is far narrower than the diameter of the aorta, and according to the law of continuity, fluid should travel faster through a narrower diameter tube, the rate is actually slower due to the overall diameter of all the combined capillaries being far greater than the diameter of the individual aorta.
The slow rate of travel through the capillary beds, which reach almost every cell in the body, assists with gas and nutrient exchange and also promotes the diffusion of fluid into the interstitial space. After the blood has passed through the capillary beds to the venules, veins, and finally to the main venae cavae, the rate of flow increases again but is still much slower than the initial rate in the aorta. Blood primarily moves in the veins by the rhythmic movement of smooth muscle in the vessel wall and by the action of the skeletal muscle as the body moves. Because most veins must move blood against the pull of gravity, blood is prevented from flowing backward in the veins by one-way valves. Because skeletal muscle contraction aids in venous blood flow, it is important to get up and move frequently after long periods of sitting so that blood will not pool in the extremities.
Blood Pressure and Velocity
The pressure of the blood flow in the body is produced by the hydrostatic pressure of the fluid (blood) against the walls of the blood vessels. Fluid will move from areas of high to low hydrostatic pressures. In the arteries, the hydrostatic pressure near the heart is very high and blood flows to the arterioles where the rate of flow is slowed by the narrow openings of the arterioles. During systole, when new blood is entering the arteries, the artery walls stretch to accommodate the increase of pressure of the extra blood; during diastole, the walls return to normal because of their elastic properties. The blood pressure of the systole phase and the diastole phase, graphed below, gives the two pressure readings for blood pressure. For example, 120/80 indicates a reading of 120 mm Hg during the systole and 80 mm Hg during diastole. Throughout the cardiac cycle, the blood continues to empty into the arterioles at a relatively even rate. This resistance to blood flow is called peripheral resistance.
Blood pressure is related to the blood velocity in the arteries and arterioles. In the capillaries and veins, the blood pressure continues to decease but velocity increases.
Exchange Across Capillaries
Proteins and other large solutes cannot leave the capillaries. The loss of the watery plasma creates a hyperosmotic solution within the capillaries, especially near the venules. This causes about 85% of the plasma that leaves the capillaries to eventually diffuses back into the capillaries near the venules. The remaining 15% of blood plasma drains out from the interstitial fluid into nearby lymphatic vessels. The fluid in the lymph is similar in composition to the interstitial fluid. The lymph fluid passes through lymph nodes before it returns to the heart via the vena cava. Lymph nodes are specialized organs that filter the lymph by percolation through a maze of connective tissue filled with white blood cells. The white blood cells remove infectious agents, such as bacteria and viruses, to “clean” the lymph before it returns to the bloodstream. After it is “cleaned,” the lymph returns to the heart by the action of smooth muscle pumping, skeletal muscle action, and one-way valves joining the returning blood near the junction of the venae cavae entering the right atrium of the heart.
Fluid from the capillaries moves into the interstitial space and lymph capillaries by diffusion down a pressure gradient and also by osmosis. Out of 7,200 liters of fluid pumped by the average heart in a day, over 1,500 liters is filtered. (credit: modification of work by NCI, NIH)
This video describes the function of the lymphatic system in conjunction with the circulatory system (stop at 5:40, when the discussion of immune function begins):