Lectures 16&17

Circulatory System
(Plumbers Guide to the Vertebrates)
Kardong Chapter 12, Hildebrand Chapter 14

Circulatory System
Distributes water, solutes (ions, nutrients, proteins), dissolved gases (O2, CO2, NH3), heat, information (from endocrine system), and cells involved in damage repair, the immune system and gas exchange. 2 parts - Blood vascular system heart, arteries, veins, capillaries, blood. - Lymphatic system lymphatic vessels, lymph nodes, lymph hearts, lymph.

Vertebrate Blood and Lymph

Plasma - composed of water, salts, nutrients, wastes, hormones, and blood proteins like albumens, fibrinogens (involved in clotting), and globulins (involved in immune response). Blood cells erythrocytes found in BV system only; contain hemoglobin; transport O2 and CO2 - leucocytes found in BV and lymph systems; diverse types, all involved in the immune response - platelets (or thrombocytes in non-mammals) involved in the clotting response.

mammals have unique, biconcave erythrocytes without nuclei

Early Development of the Circulatory System

Mesoderm tissue forms blood islands on the yolk, that coalesce into vitelline vessels containing blood.

KK 12.7

Hemopoiesis or blood cell formation

Sites of hemopoiesis change during development and have changed during evolution.
blood islands on the yolk of amniotes and embryonic blood vessels are the first sites of hemopoiesis gut and organs of the gut in embryos and fishes spleen - hemopoietic, but in mammals produces leucocytes only and stores and destroys erythrocytes. thymus - produces leucocytes, and responsible for development of the immune system in embryos. lymph nodes - leucocytes red bone marrow - ribs, centra, epiphyses in tetrapods

Components of the Circulatory System

capillaries
vein portal vein

heart

arteries
shunt

capillaries

veins

lymph vessels

KK 12.46

sinuses

Arteries, Capillaries, Veins

Pressure Change in the BV system Control of microcirculation

KK 12.2

KK 12.4, H&G 14.10

Structure of Blood Vessels

The inner or tunica intima (interna) layer includes the endothelial layer that lines the vessel and comprises capillaries. The tunica media is is a muscular layer with smooth muscles and elastic cartilage. The outer tunica adventitia (externa) is a layer of fibrous connective tissue. The relative thickness of the tunica media varies among different kinds of vessels.

KK 12.1, H&G 14.11

Circulation in the Shark

In fishes, the heart pumps blood to the gills where it is oxygenated, then the internal carotid arteries and the dorsal aorta distribute it around the body.

KK 12.11. H&G 14.12

Returning blood blood is collected via paired anterior and posterior cardinal veins (not labelled on this diagram). Tail circulation returns to the kidneys via the renal portal vein. Most vessels are paired.

KK 12.10, H&G 14.12

All vertebrate embryos, have circulatory systems with 6 aortic arches on each side of the pharynx except Agnatha (which typically have >6). Adult fish have 4 or 5 (e.g. shark). Major arteries and veins (except dorsal and ventral aorta, and vessels of the gut) are paired unlike in adult amniotes. This is an example of recapitulation.

The Primitive/Embryonic Arterial Arches

Agnatha (living and ostracoderms) have more than 6 pairs of arterial arches, but all other vertebrate embryos have 6. The adult pattern of the arches leaving the heart is created by partial deletion during development of these embryonic 6 pairs of arches.

KK 12.13,H&G 14.12

Sharks have 5 aortic arches; the first is deleted in development. The ventral aorta extends forward as the external carotid artery serving the jaw.

KK 12.14, H&G 14.13

Teleosts only have 4, and in living Sarcopterygii the 3rd and 4th no longer serve gill tissue. They are direct connections between the ventral and dorsal aorta. Note that a branch of the 6th arch serves the lung.

Amphibia
In Amphibia, arches 1&2 are deleted. The 3rd persists to carry blood to the head via the internal carotid artery. The 5th may also be deleted, but the 4th always remains as a connection between the ventral and dorsal aorta taking blood to the posterior body. The dorsal aorta is usually not continuous, but disappears between the 3rd and 4th arches. The 6th arch takes blood to the lungs. Ductus arteriosus? Carotid duct?
KK 12.14, H&G 14.13

Amniotes
As in Amphibia, aortic arches 1,2 and 5 are lost in development. The 3rd carries blood to the head, the 4th carries blood to the body, and the 6th to the lungs (pulmonary artery). In addition, the symmetry is lost. In reptiles, the left side serves the body only, while the right side serves the head and body. In birds, the redundant left 4th arch is lost completely.
In mammals, a parallel assymmetry appears, but it is the left side that becomes the systemic arch of the aorta and the right side is lost. The ductus arteriosus persists until birth. Why?

KK 12.17

Increasing asymmetry, ventral view

Common carotid arteries (Cc) are derived from the ventral aortic arches. Subclavian arteries (Sc) serve the forelimbs.

KK 12.18

Summary:
evolution of the aortic arches
The various adult patterns of the aortic arches are derived from selective deletions of the basic embryonic pattern.

Tetrapods lose arches 1,2 and 5 entirely. The 6th contributes to the pulmonary arteries.

KK 12.9, H&G 14.13

The major veins, as seen in embryos and adult fish

Embryonic and fish major veins are symmetrical. In adult fish, caudal circulation returns to the heart via the kidneys, and the artery is the renal portal vein. Where are the hepatic veins?

KK 12.21, H&G. 14.14, 14.18

Venous System of Amphibia

In Amphibia, the caudal circulation also passes through the kidneys. A new vein, the postcava takes over at least partly the circulation previously carried to the heart by the posterior cardinal veins. The postcava is derived, proximally, from the hepatic veins.

KK 12.23 d,e H&G 14.18 KK 12.23

Venous System of Amniotes

In amniotes, the venous system continues to simplify and lose its symmetry. The postcava delivers all blood from the body to the heart.

Venous System of Mammals

Mammalian venous system is simplified by loss of all traces of the embryonic paired cardinal veins, both anterior and posterior. The precava is the only vein carrying blood into the heart from the anterior, and is derived from the fusion of portions of several embryonic veins. The venous circulation does not pass through the kidney, but rather the kidney receives only arterial blood.
KK 12.23, H&G 14.18, 14.24

Early Development of the Heart

The heart develops from the splanchnic hypomere, specifically the ventral mesenteries. It begins as contractile parallel vessels that coalesce into a single linear pump. The embryonic (and adult) heart has three layers; endothelium, muscle and connective tissue. That is, it resembles an artery in its basic structure. . KK 12.8, H&G 14.1

The Embryonic Heart KK 12.24, H&G14.2

The heart of vertebrate embryos is a linear pump, sending blood forward through the ventral aorta to the branchial arches.

There are four main parts: the sinus venosus, which receives blood returning via the common cardinal veins; the atrium, which delivers blood into the main pump, the ventricle; and the conus, with distinctive conal (semilunar) valves that prevent backflow when the ventricle relaxes between beats. Note also sinuatrial and atrioventricular valves.

Fish Hearts
KK 12.27, H&G 14.3

Adult fish hearts are like embryonic hearts except that the atrium lies above the ventricle, so that the flow of blood is in an S pattern. Teleosts have an elastic conus called the bulbus arteriosus (remember the trout).
Note that the ventricle is the main pump, and is much more muscular.

Amphibians: vertebrates with different sites for gas exchange

Typical amphibian

KK 12.30, H&G Fig. 14.4

Bullfrog

Amphibian atria are divided, with blood from the lungs entering on the left, and blood entering from the sinus venosus entering on the right. The ventricle is not divided, but can keep deoxygenated blood from the body and oxygenated blood from the lungs separate when appropriate. It can direct a variable fraction of the blood entering heart to the body or to the lungs depending on whether the lungs are in use or the animal is relying on cutaneous respiration.

Lateral views of amphibian hearts and arches

In larval stages, there may be three sites for gas exchange; external gills, lungs and skin.

KK 12.31

KK 13.31

Reptile Hearts
Reptile hearts have fully divided atria and a 3-chambered ventricle. Blood returns from the lungs to the left atrium and from there goes to the deep cavum arteriosum. It passes through an interventricular canal to get to the cavum venosum, where it can be sent to the body via the left and right aortic arches. Blood returning from the body enters the right atrium. It goes over the muscular ridge separating the cavum venosum and the cavem pulmonale to the cavum pulmonale to be pumped to the lungs via the pulmonary arches. When the lungs are not being used (e.g. diving in turtles) the ridge can stop blood from crossing to the cavum pulmonale, and direct it back to the right aortic arch.

KK 12.32, H&G 14.5

Reptile Hearts 2
Taking a larger view, oxygenated blood returning to the left atrium is directed to the cavum arteriosum, under the cavum pulmonale, to avoid sending it back to the lungs. Deoxygenated blood entering the right atrium can be directed to the pulmonary arteries by letting it cross the ridge, or be directed to the right aortic arch to the head and body. The latter is appropriate when the animal is conserving O2, as the left arch mostly serves the body.

KK 12.33

KK 12.32, H&G 14.5

KK 12.35

Mammal Heart KK 12.39, H&G 14.6

The hearts of mammals and birds have fully divided atria and ventricles. Essentially they have two isolated circuits. The right ventricle takes venous blood returning from the body and sends it to the the lungs via the pulmonary arteries. The left ventricle takes blood returning from the lungs and sends it to the body. There is no mixing of the two sources of blood, so it is highly efficient but completely inflexible; appropriate for endotherms with high and relatively constant metabolic rates, and only one site for gas exchange.

Mammal Heart KK 12.42, H&G 14.6, 14.7

The right diagram illustrates the dual circuit circulation of mammals. Before birth, embryonic circulation (left) uses the ductus arteriosus and an opening in the septum between the atria to bypass the lungs.

Conservation of Heat in Endotherms

Endotherms can conserve or lose heat by altering their microcirculation patterns to heat the entire body or just the core. Limbs in cold water use countercurrent circulation to warm the cold blood returning to the body. That is, heat is lost to the incoming blood rather than the water. KK 12.43, 44

Nasal turbinates
Turbinate bones above the secondary palate support a highly vascularized area that conserves moisture, conserves heat in the cold, and also helps cool the brain in hot weather. Cooling the brain, which is very sensitive to overheating, is accomplished with the assistance of a carotid rete, where arterial blood going to the brain can be cooled by blood returning from the nasal area.

Dog

Eland (antelope)

KK 12.45

Lymph vessels and nodes carry lymph back

toward the hear via lymph nodes. Valves prevent fluid from flowing away from the heart.