21.1. Overview of the Circulatory System

For more complex organisms, diffusion is not efficient for cycling gases, nutrients, and waste effectively through the body; therefore, more complex circulatory systems evolved. Most arthropods and many mollusks have open circulatory systems. 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. Figure 21.4 illustrates the basic circulatory systems of some vertebrates: fish, amphibians, reptiles, and mammals.

As illustrated in Figure 21.4a 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 fish’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.

As shown in Figure 21.4b, 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, as shown in Figure 21.4c. 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. Together these adaptations have made crocodiles and alligators one of the most evolutionarily successful animal groups on earth.

In mammals and birds, the heart is also divided into four chambers: two atria and two ventricles, as illustrated in Figure 21.4d. The oxygenated blood is separated from the deoxygenated blood, which improves the efficiency of double circulation and is probably required for 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.

Summary

In most animals, the circulatory system is used to transport blood through the body. Some primitive animals use diffusion for the exchange of water, nutrients, and gases. However, complex organisms use the circulatory system to carry gases, nutrients, and waste through the body. Circulatory systems may be open (mixed with the interstitial fluid) or closed (separated from the interstitial fluid). 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 adaptions during evolution and associated differences in anatomy. Fish have a two-chambered heart with unidirectional circulation. Amphibians have a three-chambered heart, which has some mixing of the blood, and they have double circulation. Most non-avian reptiles have a three-chambered heart, but have little mixing of the blood; they have double circulation. Mammals and birds have a four-chambered heart with no mixing of the blood and double circulation.