he circulatory (or cardiovascular) system is responsible for transporting nutrients, gases, hormones and metabolic wastes to and from

the individual cells of an organism. Mammals are far too large for all of their individual cells to exchange nutrients, wastes and gases with the external world by simple diffusion. Most cells are buried too deep inside the body for this method to be effective. Thus, some system must be in place to efficiently exchange these products between the outside world and every cell in the organism's body. For this reason, the circulatory system is a highly-branched network of vessels that spreads throughout the entire organism. In general the circulatory system represents a series of vessels that diverge from the heart (arteries) to supply blood to the tissues and a confluence of vessels draining blood from the tissues and returning it to the heart (veins). Despite the extensive network of arteries and veins throughout the body, no actual exchange of water, nutrients, wastes or gases occurs in arteries or veins. Their walls are too thick to permit diffusion. Extensive networks of capillary beds connecting branches of arteries and veins exist throughout the body to transfer these dissolved substances between the bloodstream and the tissues. To simplify the identification of the numerous arteries and veins, there are two general principles you should remember: (1) arteries and veins tend to be paired, especially when the organs they supply or drain are paired, and (2) a continuous vessel often undergoes several name changes along its length as it passes through different body regions. Therefore to successfully identify arteries and veins it is necessary to trace them along their entire length (typically from the heart outward).

outward in all directions (Figure 5.2). For the moment, we will concentrate only on the heart and the major vessels that originate from it. Later we will trace the paths of these major vessels as they diverge throughout the body. Mammals possess a four-chambered heart that delivers blood through two major circulatory pathways-the pulmonary circuit (from the heart to the lungs and back) and the systemic circuit (from the heart through the rest of the body and back). A hallmark of the mammalian heart is that its internal design keeps the blood from these two circuits entirely separate, thus keeping oxygen-depleted blood from mixing with oxygen-rich blood.
NOTE: Throughout this chapter, we use the terms atrium and auricle to refer to slightly different regions of the heart. The term auricle is used to describe the small, outer, flap-like region that covers a portion of the atrial chamber, while the term atrium is used to refer to the entire open space (or actual chamber) inside that collects the blood. The reason for this distinction is that part of each atrial chamber extends well beyond the boundaries of each flap-like auricle. This will be more evident when you view the interior of the heart later.

THORACIC CAVITY AND NECK REGION

Notice the thin pericardial membrane surrounding the heart (Figure S.lA). This protective sac contains a small amount of lubricating fluid to protect the heart and cushion its movements. A portion of the relatively large thymus sits on the outside of the pericardial membrane and partially obscures the heart from view. The thymus should not be confused with neighboring lung tissue or with the atria of the heart (which lie inside the pericardium). Make a mental note of the position, color and consistency of the thymus. We will return to the thymus along with the other endocrine glands in Chapter 9.

The Heart: External Anatomy

The driving force behind the entire circulatory system of mammals is the heavily muscled heart. Numerous vessels emanate from the cranial aspect of the heart and radiate

Identify the four chambers of the heart. Caudally there are two large, thick-walled ventricles, the right ventricle and the left ventricle (Figure S.lB). These chambers pump blood out of the heart to the lungs and to the rest of the body, respectively. Cranial to the ventricles and somewhat darker in color are the right and left auricles. Chambers within the right and left auricles receive blood from the body and the lungs, respectively, and pass it to the ventricles. Running along the surface of the heart itself, the small coronary arteries (Figure S.2A) should be evident. These vessels supply blood to the heart muscle, insuring that it too receives nutrients and oxygen. Notice the large veins entering the heart on the right side. These are the cranial and caudal vena cavae (Figure S.2A). They bring deoxygenated blood to the right atrium from the upper and lower portions of the body. On the dorsal surface of the heart, adjacent to the juncture of the vena cavae and the right atrium, there is a small sac-like region of the heart known as the coronary sinus (Figure S.2E). This sinus is responsible for returning deoxygenated blood from the wall of the heart to the right atrium. On the ventral surface of the heart, locate the large pulmonary artery (Figure S.2D) emanating from the right ventricle. In the adult, this artery channels blood from the right ventricle through the right and left pulmonary arteries to the lungs. Follow the pulmonary artery behind the heart and see where it branches into the right and left pulmonary artery. Notice that at the base of the pulmonary artery there is a connection between it and the aorta (the large artery leaving the cranial aspect of the left ventricle). This connection is called the ductus arteriosus (Figure 5.2C), a short, temporary linkage found only in the fetus. Lying adjacent to the pulmonary arteries are the pulmonary veins, the vessels that, in the adult, return oxygenated blood to the left atrium of the heart.

Thyroid gland
Artery

Vein

Pericardium Lung

Diaphragm

Cranial vena cava Aortic arch Pulmonary artery Left auricle Left lung Coronary and vein artery

Right auricle

Right ventricle

Left ventricle

m
II FIGURE 5.1 Heart surrounded by perieardial membrane (a), heart with perieardial membrane removed (b), and inset

showing histology oj an artery and vein (e).

Vagus nerve Right costocervical vein Cranial vena cava Right auricle Branches of the coronary artery Right ventricle Caudal vena cava Cranial vena cava Right auricle Pulmonary artery Right ventricle Heart reflected laterally Left ventricle Left phrenic nerve Brachiocephalic trunk Left subclavian artery Ductus arteriosus Left coronary artery Left auricle reflected

Pulmonary vein and artery Phrenic nerve Accessory lobe of right lung Diaphragm

II
Right internal thoracic vein (cut) Cranial vena cava Aorta Ductus arteriosus Pulmonary artery Coronary vessels Left auricle Left lung Phrenic nerve Left costocervical artery and vein Left internal thoracic artery Brachlocephalic trunk Left subclavian artery Accessory lobe of the right lung

Cranial vena cava Pulmonary artery

Left costocervical vein Vagus nerve Aortic arch Phrenic nerve Pulmonary Bronchus veins

m
Heart reflected cranially Left auricle Right phrenic nerve Coronary Right lung Caudal vena cava sinus vein Left pulmonary Accessory lobe of the right lung

Phrenic nerve terminating on the diaphragm

III

FIGURE 5.2 Composite photograph oj heart anatomy, cranial and caudal vena cavae {al, right atrium and coronary arteries {bl, aorta, pulmonary trunk, and ductus arteriosus {cl, pulmonary arteries and veins {dl and coronary sinus and caudal vena cava {el.

Fetal v. Adult Circulation

There are three major differences in the circulatory system of a fetal and adult pig (Figure 5.3). The most obvious is the connection between the fetus and the mother through the umbilical cord (Figure 5.3C). This collection of tubes allows oxygen and nutrients to pass from the mother to the fetus while transporting carbon dioxide and metabolic wastes away from the fetus. The other two major differences lie in the structure of the heart. Since the fetus is not breathing with its lungs, the lungs do not oxygenate blood that passes through them. In fact, the most oxygen-rich

Cranial vena cava Ductus arteriosus

Cranial vena cava Toand from upper extremity

Caudal

At birth Umbilical arteries Allantoic stalk

Umbilical vein

To and from lower extremity FIGURE

Umbilical vein

Urogenital

opening

with inset photograph of umbilical cord (c).

5.3 Schematic illustrations of the fetal circulatory pathway (a) and the comparative pathway after birth (b),

blood in the fetal circulatory system is that returning to the fetus through the umbilical vein. As a result, only a small fraction of the blood leaving the right ventricle travels through the pulmonary arteries to the lungs. The majority is redirected through the ductus arteriosus, a connection between the pulmonary artery and the aorta that channels blood into the aorta (Figure 5.2C). Another structure inside the heart of the fetus, called the foramen ovale, also aids in rerouting blood to bypass the lungs. This opening in the septum between the right and left atria allows blood passing into the right atrium to be channeled into the left atrium and away from the lungs. Both of these adaptations

ensure that the majority of oxygenated blood arriving via the umbilical vein is passed through the fetal circulatory system via the aorta, while permitting enough blood to reach the lungs to allow the lung tissue to develop.

Veins oj the Thoracic Region

The largest veins in the thoracic region are the cranial vena cava and caudal vena cava, which converge at the entrance to the right atrium (Figures 5.4-5.5). In the adult, these two thin-walled veins bring deoxygenated blood back to the heart from all parts of the body. Trace one cranial vena

Internal jugular vein

External jugular vein Cephalic vein (cut)

Thyroid gland Axillary artery and vein

Right brachlocephallc vein Internal thoracic vein

Subclavian vein

Costocervical trunk

Cranial vena cava

Right ventricle

Left ventricle

FIGURE

5.4 Veins oj the thoracic region.

cava cranially to its first major branch. This short branch, known as the brachia cephalic vein (or trunk) represents the confluence of four veins: the internal jugular vein, the external jugular vein, the cephalic vein, and the subclavian vein. Remember, blood is flowing back toward the heart through these vessels. Identify the internal thoracic vein leading from the arm pit toward the vena cava and heart at a ninety degree angle. The left and right axillary veins also lead toward the vena cava at ninety degree angles and bring blood from the forelimbs of the pig. The subscapular vein leading from the arm pit and the axillary vein come

together to form the subclavian vein which dumps blood directly into the cranial vena cava. The third vein returning blood from each forelimb is the cephalic vein, the most cranial of the three. The external jugular veins lead down into the vena cava from the neck region, along with the internal jugular veins running medially alongside the trachea from the head toward the heart. Follow the external jugular vein cranially to the point where it bifurcates into the linguofacial vein and the maxillary vein. These veins return blood from the temporal portion of the face and lower jaw, respectively.

Linguofacial

vein

Maxillary vein

External jugular vein Right subclavian vein

Internal jugular vein

Cephalic vein

Subscapular

Long thoracic vessels

Right brachiocephalic

vein Costocervical vein

Cranial vena cava Caudal vena cava

Left ventricle

Diaphragm

FIGURE

5.5 Veins oj the thoracic region.

Arteries oj the Thoracic Region

Earlier, you identified the large aorta emanating from the cranial aspect of the left ventricle. Now, trace the aortic arch as it curves caudally (Figure 5.6). The first major branch off the aorta is the brachiocephalic trunk, which immediately splits into the right subclavian artery (which carries blood to the right forelimb and upper potion of the body) and the carotid trunk, off which the common carotid arteries branch

(which carry blood to the head and brain). Follow the common carotid cranially to the point where it bifurcates into the external carotid artery (which runs along the ventral side of the masseter) and the internal carotid artery (which embeds underneath the masseter muscle) (Figures 5.7-5.8). The axillary artery is a continuation of the right subclavian artery which carries blood into the armpit and shoulder region. A small artery may be seen branching off the axillary artery and leading caudally toward the ribs. This is the internal thoracic artery. The second major branch off the aorta is the left subclavian artery which carries blood to the left forelimb and left portion of the upper body. Locate the other axillary artery, the continuation of the left subclavian artery. The aorta continues caudally along the dorsal body wall and passes through the diaphragm into the abdominal cavity. At this point it is commonly called the dorsal aorta. You will have to move the lobes of the left lung toward the center of the pig to view this. (DO NOT remove the lungs yet!)

Thyroid gland

Esophagus Right vagus nerve Left vagus nerve

Right phrenic nerve Right subclavian artery

Common

carotid artery

Trachea Brachiocephalic trunk Left subclavian artery Aortic arch Ductus arteriosus

Cranial vena cava (cut) Right auricle

Pulmonary

artery

Left auricle

FIGURE

5.6 Major arteries oj the thoracic region (veins have been removed for clarity).

Internal carotid artery

External carotid artery

Arterial branches to the thyroid gland Right subclavian artery Thyrocervical trunk Axillary artery External thoracic artery
---,L-l,l':"""'_''+--II---

Internal thoracic artery --_J.--H~r--Z Brachiocephalic trunk Cranial vena cava (cut)

Left subclavian artery Costocervical artery Arch of aorta

Left coronary artery Caudal vena cava

Pulmonary artery Left ventricle

Diaphragm

FIGURE

5.8 Arteries oj the thoracic region (veins have been omitted ior clarity).

The Heart: Internal Anatomy

juncture of the right atrium and right ventricle is the tricuspid valve. As the right ventricle contracts and pushes blood out to the lungs, the blood is forced back up against the tricuspid valve, closing the leaflets of the valve and preventing retrograde flow into the right atrium. Upon entering the pulmonary trunk, the blood also passes through the pulmonary semilunar valve, which prevents backflow into the right ventricle. The blood returns from the lungs into the left atrium via the pulmonary veins and then flows into the left ventricle through the bicuspid (or mitral) valve. Blood leaving the left ventricle into the aorta is regulated by the aortic semilunar valve. The valves of the heart are prevented from being pushed too far backward (a condition known as "prolapse") by small stringlike attachments called chordae tendineae. You may be able to see these structures on your frontal section of the heart.

Notice that inside the chambers of the heart there are valves to prevent blood from flowing backwards (Figures 5.9-5.10). As blood enters the right atrium, it immediately flows into the right ventricle. Very little blood is actually pumped by the right atrium into the right ventricle. At the

Left common carotid artery

Internal jugular vein (cut)

Vagus nerve

Right common carotid artery Right thoracocervical artery

Left thoracic cervical trunk

Left axillary artery and brachial plexus

Right axillary artery and nerve

Right external thoracic artery Right internal thoracic artery Right subclavian artery Brachiocephalic trunk FIGURE

Left subclavian artery

Left ventricle

5.7 Arteries oj the thoracic region (veins have been removed jor clarity).

----

Left subclavian artery Aorta

Cranial vena cava Right pulmonary artery Right atrium

Cranial vena cava Right auricle Pulmonary artery

-----1

veins Right ventricle Coronary vessels --Caudal vena cava

© Michael Schenk

Right pulmonary vein Caudal vena cava

Left auricle

Coronary

sinus

Left ventricle --Right ventricle

OJ

Brachiocephalic

artery

Opening

to coronary

artery

Pulmonary

trunk

Ductus arteriosus at junction of aorta and pulmonary trunk Left atrium

Semilunar

valve

Azygos vein

Chordae tendinae ---Left ventricle

Papillary muscle Left ventricle

heart (c and dJ.

FIGURE

5.9 Illustrations of heart depicting ventrolateral view (aJ, dorsal view (bJ and interior views through frontal plane of

Left subclavian Brachiocephalic trunk Cranial vena cava Pulmonary artery Right auricle Left auricle

Costocervical

Aorta Cranial vena cava Azygos vein Right pulmonary artery Left auricle Pulmonary veins Coronary sinus Right atrium Caudal vena cava Right coronary artery and vein

Right ventricle Left coronary artery and vein Left ventricle

Left ventricle Aorta Brachiocephalic trunk Left subclavian artery Pulmonary artery

Pulmonary

artery

Ductus arteriosus Opening coronary to left artery Right atrium Left coronary Bicuspid valve artery

Tricuspid valve

Right ventricle Right ventricle

of heart (e).

FIGURE

5.10 Photographs of heart depicting ventral view (a), dorsal view (b) and interior views through frontal plane

ABDOMINAL

CAVITY

Hepatic Portal System

Follow the caudal vena cava from the heart through the diaphragm and liver toward the stomach. Notice how it passes directly through the diaphragm and through the center of the lobes of the liver. Situated below the liver, among the intestines, pancreas, spleen and stomach there is a

unique system of veins called the hepatic portal system (Figures 5.11-5.12). Portal systems are found in many different parts of the body in mammals. Their purpose is to shunt blood between the capillary beds of certain organs before allowing it to pass along the rest of the body. Portal systems consist entirely of veins that reroute blood between organs without passing through the heart.

Hepatic portal vein

Left gastroepiploic Gallbladder ---

vein

Cystic duct

Spleen

Common

bile duct

Splenic vein

Hepatic vein

Right gastroepiploic

vein

Ductus venosus

Pancreas ---+.-1-'0 (right lobe)

Caudal vena cava

Umbilical vein

,.----

Hepatic portal vein

---

Small intestine (ileum)

Portal sinus

FIGURE

5.11 Illustration oj hepatic portal system depicting associated veins and organs.

Common Hepatic portal vein

bile duct

Mesenteric vein

Gastrosplenic

vein

Aorta

Vagus nerve Hepatic artery Umbilical vein Gallbladder Cystic artery over cystic duct Left gastric artery Celiac artery Stomach Hepatic portal vein Pancreas Small intestine (duodenum)

Cranial mesenteric artery and mesenteric vein Cecum

Small intestine (jejunum)

Small intestine (ileum)

m
FIGURE

5.12 Hepatic portal system with liver partially removed and duodenum displaced.

In the case of the hepatic portal system, blood flows from the capillary beds of the small and large intestines, the spleen, the pancreas and the stomach into the hepatic portal vein (Figure 5.12) and then into the capillary beds of the liver, before entering the caudal vena cava. This extra step allows blood from the stomach and intestines that contains large amounts of sugars and possibly toxins to be filtered by the liver before the blood is sent to the rest of the body. Also, hormones produced by the pancreas (e.g., insulin, glucagon, somatostatin) can be directed to their target organ, the liver, without the delay and diluting effect of traveling through the

entire circulatory system first. Depending on the type and amount of hormone released by the pancreas, the liver may store the sugar (as glycogen) or release it into the bloodstream immediately. Through this regulatory mechanism, the liver maintains nearly constant blood glucose levels.

Arteries and Veins oj the Abdominal Region

As the aorta passes caudally through the abdominal region it makes several more branches (Figures 5.13-5.15). Locate the celiac artery, a small branch from the aorta to the

Umbilical vein (cut)

-+--"1--- Renal artery and vein

Genital vessels (spermatic-male, ovarian-female)

Caudal mesenteric artery

Median sacral artery and vein

Deep circumflex iliac artery and vein

External iliac artery and

© Michael Schenk

Umbilical arteries FIGURE

Urinary bladder

5.13 Arterial supply and venous return oj organs in the abdominal cavity and the lower extremities.

stomach, pancreas and spleen. Next find the cranial mesenteric artery (Figure 5.14A), which has branches that supply the jejunum, ileum and colon. Embedded in the intestinal mesentery are the arterial arcades (Figure 5.14B), numerous branches of the mesenteric artery that provide nutrients and oxygen to the tissues of the intestinal tract. Further caudally, two short branches of the dorsal aorta lead into

the kidneys. These are the renal arteries. Lying next to the renal arteries, the thinner-walled renal veins are present. These vessels collect filtered blood from the kidneys. The caudal mesenteric artery is a single vessel that supplies blood to the colon and rectum (Figure 5.15A). Caudal to this vessel, the genital arteries (Figure 5.15B) are visible branching off the dorsal aorta and leading to the ovaries or

Spiral colon Pancreas Adrenal gland

Cranial mesenteric artery and mesenteric vein Renal artery and vein Small intestine (ileum)

Caudal vena cava

Spiral colon

Cranial mesenteric artery

Mesenteric lymph nodes

Small intestine (jejunum) Small intestine (ileum)

Arterial arcades of the mesentery (from branches of cranial mesenteric artery)

m
II FIGURE 5.14 Cranial mesenteric artery (a), arterial arcades oj the cranial mesenteric artery (b), and cranial

mesenteric, celiac, and caudal mesenteric arteries and their branches (c),

testes, depending upon the sex of the pig. The genital veins run alongside the corresponding arteries but lead back into the caudal vena cava. If you follow the dorsal aorta caudally to the point where it branches into each hindlimb, you should be able to identify several more arteries and veins. Locate the external iliac arteries and external iliac veins leading into the upper thigh of each hindlimb. These

vessels supply and receive blood from the legs. Further along, as they pass from the abdomen into the hindlimb, they become the femoral arteries and femoral veins and then branch into the deep femoral arteries and deep femoral veins. Finally, locate the internal iliac arteries and internal iliac veins, lying dorsal to the colon.

Common bile duct

Hepatic artery

Umbilical vein (cut) Cystic artery on cystic duct Gallbladder Hepatic portal vein Small intestine (duodenum) Gastroduodenal artery and vein Right gastric artery and vein Hepatic artery Splenic artery and gastrosplenic vein (cut) Celiac artery Cranial mesenteric artery Left adrenal artery Renal vein Renal artery

Mesenteric

vein

Branches of the cranial mesenteric artery Caudal vena cava Small intestine: jejunum ileum

Aorta Caudal mesenteric artery

Umbilical artery

External iliac ---artery and vein

Renal vein Renal artery Caudal Kidney

Colon

Ureter Aorta

Branches of caudal mesenteric artery supplying distal colon and rectum

Caudal mesenteric artery Ovarian artery

External iliac artery

Ureter (cut) Aorta Caudal vena cava Testicular artery and vein Umbilical arteries Caudal mesenteric artery (cut) Deep circumflex artery and vein Common Median sacral artery iliac vein

Deep femoral artery Femoral artery

External iliac artery and vein Internal iliac artery and vein

Femoral artery, vein and nerve Deep femoral artery and vein Pelvis (cut) and colon removed

removed for clarity.

III

FIGURE

5.15 Caudal mesenteric artery (a) and genital, iliac, and femoral arteries {b)j rectum cut and digestive tract

Umbilical Cord

storage. Thus the urinary bladder remains connected to the umbilical cord until the fetus is born (Figure 5.17). After birth, the connection to the umbilical cord deteriorates and the urinary bladder drains into the urethra.

In placental mammals, the unborn young are attached to the placenta of the mother via the umbilical cord (Figure 5.16). This cord serves as the lifeline for the fetus, transporting nutrients and oxygen to the growing fetus, and providing a channel for carbon dioxide and excess metabolic wastes to be eliminated from the fetus. Identify the single umbilical vein. Unlike most veins, this vessel actually carries oxygen- and nutrient-rich blood to the fetus from the fetal side of the placenta. The maternal blood is restricted to the maternal side of the placenta and never mixes with fetal blood under normal circumstances. Exchange of gases, nutrients and wastes is accomplished by diffusion across the placental barrier. You should see two smaller umbilical arteries (Figures 5.16-5.17). These carry blood from the fetus to the placenta. While the fetus remains in its mother's uterus, metabolic waste (urine) that collects in the bladder passes through the allantoic duct and into the allantois for

Umbilical arteries FIGURE

Urogenital opening

5.16 Distal end oj umbilical cord.

Umbilical artery

Branches of umbilical artery supplying the urinary bladder

Urinary bladder

Ureter

III FIGURE 5.17 Close-up oj umbilical arteries and urinary bladder.

The Spleen
The spleen is a vascular, ductless organ that plays a critical role in the circulatory system of vertebrates (Figure 5.18). Since mammalian red blood cells do not contain nuclei, they cannot undergo cell division and thus have a finite life span. New red blood cells are continuously produced in the bone marrow and delivered to the spleen. The spleen also manufactures white blood cells (lymphocytes) to fend off diseases, and destroys and recycles worn-out blood cells. The spleen stores these cells along with excess blood plasma to be released into the bloodstream as needed. Through this mechanism the spleen regulates the body's total blood volume and the concentration of red blood cells in the blood.

Splenic artery and vein Stomach

Small intestine

III FIGURE 5.18 Spleen (displaced to the side) with accompanying histology photograph.

Spleen.

250X

THE SHEEP HEART

NOTE: Due to the popularity of the sheep heart in many laboratory courses, the following section will concentrate on the anatomy of the sheep heart as a model for studying the "typical" mammalian heart. Much of this information was covered previously with the fetal pig heart. Check with your instructor to see if you will be completing this section and to determine what level of depth you will be required to know. Identify the four chambers of the heart. Caudally there are two large, thick-walled ventricles, the right ventricle and the left ventricle (Figure 5.19). These chambers pump blood out of the heart to the lungs and to the rest of the body, respectively. In the sheep heart, there is a superficial landmark separating these two chambers known as the interventricular groove that runs obliquely down the ventral surface of the heart toward the apex, but runs more longitudinally on the dorsal surface of the heart. Cranial to the ventricles and somewhat darker in color are the right and left auricles. Chambers within the right and left auricles receive blood from the body and the lungs, respectively, and pass it to the ventricles. Running along the surface of the heart itself are the small coronary arteries and veins. The coronary arteries supply blood to the heart muscle, ensuring that it too receives nutrients and oxygen to maintain an energy supply to support its continuous, methodical beating throughout the entire life of the animal. In the sheep, the coronary vessels are typically buried under dense fat on the surface of the heart.

pulmonary artery to the aorta during fetal development. This band of solid connective tissue now joining these two vessels is known as the ligamentum arteriosum. In the sheep heart, the large brachiocephalic artery will also be visible as the first branch off the aorta. On the dorsal surface of the heart you should be able to identify the pulmonary veins leading back to the left auricle.

Notice remnants of the large veins entering the heart on the right side. These are the cranial and caudal vena cavae (Figure 5.19). They bring deoxygenated blood to the right atrium from the upper and lower portions of the body. The most visible artery from the ventral surface leaving the heart is the large pulmonary artery (pulmonary trunk) emanating from the right ventricle. This artery channels blood from the right ventricle through the right and left pulmonary arteries to the lungs. Also notice the large, thiclcwalled aorta (aortic arch) leaving the heart from the cranial aspect of the left ventricle. The aorta and pulmonary artery are connected externally for a short distance by remnant tissue of the ductus arteriosus that diverted blood from the

Notice that the atrial chambers extend well beyond the boundaries of each auricle (as mentioned earlier) and that the walls of the atria are much thinner than the walls of the ventricles. Since the ventricles are responsible for pumping blood much greater distances, they have evolved into heavily muscularized chambers capable of generating massive pressure to force blood out of the heart and through the body. Notice that inside the chambers of the heart there are valves to prevent blood from flowing backwards (Figure 5.20). As blood enters the right atrium, it immediately flows into the right ventricle. Very little blood is actually pumped by the right atrium into the right ventricle. At the juncture of the right atrium and right ventricle there is a tricuspid valve. As the right ventricle contracts and pushes blood out to the lungs, some blood is forced back up against the tricuspid valve, slamming its leaflets shut and preventing retrograde flow into the right atrium. Upon entering the pulmonary trunk, the blood also passes through the pulmonary semilunar valve, which prevents backflow into the right ventricle as it relaxes and receives more blood from the right atrium. Fully oxygenated blood returns from the lungs into the left atrium via the pulmonary veins and then flows into the left ventricle through the bicuspid valve (or mitral valve). Blood leaving the left ventricle into the aorta passes through the aortic semilunar valve, another valve to prevent baclcflow as this ventricle relaxes. The bicuspid and tricuspid valves are prevented from being pushed too far backward (a condition known as "prolapse") by small stringlike attachments of connective tissue called chordae tendineae which have small muscular attachments (papillary muscles) to the inner wall of the heart (Figure 5.20).

Brachlocephallc

artery

Aorta

Cranial vena cava Right auricle of right atrium

Ligamentum arteriosum Pulmonary artery Left auricle of left atrium

Right ventricle

Left ventricle Interventricular groove

Apex

Aorta Pulmonary artery

Brachlocephallc

artery

Cranial vena cava

Caudal vena cava Pulmonary veins Left auricle Left atrium Atrioventricular groove Right auricle Right atrium Pulmonary vein

Left ventricle

Right ventricle Interventricular groove

FIGURE 5.19 Ventral view (a) and dorsal view (b) oj the sheep heart.

Pulmonary Aorta Cranial vena cava

artery

To left atrium Bicuspid valve Chordae tendineae

Right atrium Tricuspid valve

Papillary muscles

Opening Right ventricle Interventricular septum Opening

of right coronary of brachlocephallc

artery artery

Opening of brachlocephallc artery Pulmonary artery Opening of cranial vena cava Coronary vessel

Opening of coronary sinus Bicuspid valve Right atrium Left ventricle Tricuspid valve Right ventricle Interventricular septum

Aortic semilunar valve Opening of left coronary artery

5.20 Dorsal half of frontal section (a) and ventral half of frontal section (b) of the sheep heart with inset (c) depicting openings to coronary arteries.
FIGURE