CAMPBELL BIOLOGY TENTH EDITION Reece Urry Cain Wasserman Minorsky Jackson 42 Circulation and Gas Exchange Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick
1. Circulation in the Animal Kingdom
Circulatory systems link exchange surfaces with cells throughout the body Small molecules can move between cells and their surroundings by diffusion Diffusion is only efficient over small distances because the time it takes to diffuse is proportional to the square of the distance In small or thin animals, cells can exchange materials directly with the surrounding medium In most animals, cells exchange materials with the environment via a fluid-filled circulatory system
Gastrovascular Cavities A gastrovascular cavity functions in both digestion and distribution of substances throughout the body The gastrovascular cavity is only two cells thick Flatworms have a gastrovascular cavity and a flat body that minimizes diffusion distances Mouth Radial canals Circular canal (a) The moon jelly Aurelia, a cnidarian 2.5 cm Mouth Gastrovascular 1 mm Pharynx cavity (b) The planarian Dugesia, a flatworm
Open and Closed Circulatory Systems A circulatory system has A circulatory fluid A set of interconnecting vessels A muscular pump, the heart The circulatory system connects the fluid that surrounds cells with the organs that exchange gases, absorb nutrients, and dispose of wastes Circulatory systems can be open or closed and vary in the number of circuits in the body
Figure 42.3 In insects, other arthropods, and some molluscs, hemolymph bathes the organs directly in an open circulatory system (a) An open circulatory system Heart (b) A closed circulatory system Heart Hemolymph in sinuses Pores Small branch vessels in each organ Dorsal vessel (main heart) Auxiliary hearts Interstitial fluid Blood Tubular heart Ventral vessels In a closed circulatory system, blood is confined to vessels and is distinct from the interstitial fluid Annelids, cephalopods, and vertebrates have closed circulatory systems
Organization of Vertebrate Circulatory Systems Humans and other vertebrates have a closed circulatory system called the cardiovascular system The three main types of blood vessels are arteries, veins, and capillaries Arteries branch into arterioles and carry blood away from the heart to capillaries Networks of capillaries called capillary beds are the sites of chemical exchange between the blood and interstitial fluid Venules converge into veins and return blood from capillaries to the heart
Epithelium Capillary Basal lamina Valve Epithelium Smooth muscle Connective tissue Epithelium Smooth muscle Connective tissue Artery Vein Arteriole Venule Arteries and veins are distinguished by the direction of blood flow, not by O 2 content
Single Circulation (a) Single circulation: fish Bony fishes, rays, and sharks have single circulation with a twochambered heart In single circulation, blood leaving the heart passes through two capillary beds before returning Artery Heart: Atrium (A) Ventricle (V) Vein Key Oxygen-rich blood Oxygen-poor blood Gill capillaries Body capillaries
Double Circulation (c) Double circulation: mammal Amphibians, reptiles, and mammals have double circulation Oxygen-poor and oxygen-rich blood are pumped separately from the right and left sides of the heart Key Pulmonary circuit A V Right A V Left Systemic circuit Oxygen-rich blood Oxygen-poor blood Lung capillaries Systemic capillaries
Evolutionary Variation in Double Circulation Frogs and other amphibians have a threechambered heart: two atria and one ventricle The ventricle pumps blood into a forked artery that splits the ventricle s output into the pulmocutaneous circuit and the systemic circuit When underwater, blood flow to the lungs is nearly shut off (b) Double circulation: amphibian Atrium (A) Key Pulmocutaneous circuit Right Left Ventricle (V) Atrium (A) Systemic circuit Oxygen-rich blood Oxygen-poor blood Lung and skin capillaries Systemic capillaries
2. Mammalian Circulation
Mammalian Circulation Blood begins its flow with the right ventricle pumping blood to the lungs via the pulmonary arteries In the lungs, the blood loads O 2 and unloads CO 2 Oxygen-rich blood from the lungs enters the heart at the left atrium via the pulmonary veins It is pumped through the aorta to the body tissues by the left ventricle
Figure 42.5 Superior vena cava Pulmonary artery Capillaries of right lung 9 Aorta 6 7 Capillaries of head and forelimbs Pulmonary artery Capillaries of left lung 3 11 2 4 3 Pulmonary vein Right atrium Right ventricle 10 1 5 Pulmonary vein Left atrium Left ventricle Inferior vena cava Aorta 8 Capillaries of abdominal organs and hind limbs
The aorta provides blood to the heart through the coronary arteries Blood returns to the heart through the superior vena cava (blood from head, neck, and forelimbs) and inferior vena cava (blood from trunk and hind limbs) The superior vena cava and inferior vena cava flow into the right atrium
The Mammalian Heart: A Closer Look The two atria have relatively thin walls and serve as collection chambers for blood returning to the heart The ventricles have thicker walls and contract much more forcefully
Figure 42.6 Pulmonary artery Right atrium Aorta Pulmonary artery Left atrium Semilunar valve Semilunar valve Atrioventricular (AV) valve Atrioventricular (AV) valve Right ventricle Left ventricle
The heart contracts and relaxes in a rhythmic cycle called the cardiac cycle The contraction, or pumping, phase is called systole The relaxation, or filling, phase is called diastole 1 Atrial and ventricular diastole 0.4 sec 2 Atrial systole and ventricular diastole 0.1 sec 0.3 sec 3 Ventricular systole and atrial diastole
The heart rate, also called the pulse, is the number of beats per minute The stroke volume is the amount of blood pumped in a single contraction The cardiac output is the volume of blood pumped into the systemic circulation per minute and depends on both the heart rate and stroke volume
Four valves prevent backflow of blood in the heart The atrioventricular (AV) valves separate each atrium and ventricle The semilunar valves control blood flow to the aorta and the pulmonary artery The lub-dup sound of a heart beat is caused by the recoil of blood against the AV valves (lub) then against the semilunar (dup) valves Backflow of blood through a defective valve causes a heart murmur
Maintaining the Heart s Rhythmic Beat Some cardiac muscle cells are autorhythmic, meaning they contract without any signal from the nervous system The sinoatrial (SA) node, or pacemaker, set the rate and timing at which cardiac muscle cells contract Impulses that travel during the cardiac cycle can be recorded as an electrocardiogram (ECG or EKG) Impulses from the SA node travel to the atrioventricular (AV) node At the AV node, the impulses are delayed and then travel to the Purkinje fibers that make the ventricles contract
Figure 42.8-4 1 Signals (yellow) from SA node spread through atria. 2 Signals are delayed at AV node. 3 Bundle branches pass signals to heart apex. 4 Signals spread throughout ventricles. SA node (pacemaker) AV node Bundle branches Heart apex Purkinje fibers ECG
The pacemaker is regulated by two portions of the nervous system: the sympathetic and parasympathetic divisions The sympathetic division speeds up the pacemaker The parasympathetic division slows down the pacemaker The pacemaker is also regulated by hormones and temperature
3. Blood Flow & Blood Pressure
Blood Vessel Structure and Function A vessel s cavity is called the central lumen The epithelial layer that lines blood vessels is called the endothelium The endothelium is smooth and minimizes resistance
Capillaries have thin walls, the endothelium plus its basal lamina, to facilitate the exchange of materials and are barely wider than a red blood cell Arteries and veins have an endothelium, smooth muscle, and connective tissue Arteries have thicker walls than veins to accommodate the high pressure of blood pumped from the heart In the thinner-walled veins, blood flows back to the heart mainly as a result of muscle action, and backflow is prevented by valves
LM 15 µm LM Figure 42.9 Artery Vein Artery Endothelium Smooth muscle Red blood cells 100 µm Connective tissue Capillary Basal lamina Endothelium Smooth muscle Connective tissue Valve Vein Arteriole Venule Red blood cell Capillary
Blood Flow Velocity Physical laws governing movement of fluids through pipes affect blood flow and blood pressure Velocity of blood flow is slowest in the capillary beds, as a result of the high resistance and large total cross-sectional area Blood flow in capillaries is necessarily slow for exchange of materials
Figure 42.10 Area (cm 2 ) Velocity (cm/sec) Pressure (mm Hg) Aorta Arteries Arterioles Capillaries Venules Veins Venae cavae 5,000 4,000 3,000 2,000 1,000 0 50 40 30 20 10 0 120 100 80 60 40 20 0 Diastolic pressure Systolic pressure
Blood Pressure Blood flows from areas of higher pressure to areas of lower pressure Blood pressure is the pressure that blood exerts in all directions, including against the walls of blood vessels The recoil of elastic arterial walls plays a role in maintaining blood pressure The resistance to blood flow in the narrow diameters of tiny capillaries and arterioles dissipates much of the pressure
Changes in Blood Pressure During the Cardiac Cycle Systolic pressure is the pressure in the arteries during ventricular systole; it is the highest pressure in the arteries Diastolic pressure is the pressure in the arteries during diastole; it is lower than systolic pressure A pulse is the rhythmic bulging of artery walls with each heartbeat
Regulation of Blood Pressure Vasoconstriction is the contraction of smooth muscle in arteriole walls; it increases blood pressure Vasodilation is the relaxation of smooth muscles in the arterioles; it causes blood pressure to fall Nitric oxide is a major inducer of vasodilation The peptide endothelin is a strong inducer of vasoconstriction
Figure 42.11 Measuring Blood Pressure Pressure in cuff greater than 120 mm Hg Pressure in cuff drops below 120 mm Hg Pressure in cuff below 70 mm Hg Cuff inflated with air 120 120 70 Artery Sounds closed audible in 1 2 stethoscope 3 Sounds stop
Fainting is caused by inadequate blood flow to the head Animals with long necks require a very high systolic pressure to pump blood a great distance against gravity Blood is moved through veins by smooth muscle contraction, skeletal muscle contraction, and expansion of the vena cava with inhalation Direction of blood flow in vein (toward heart) Valve (open) Skeletal muscle Valve (closed) One-way valves in veins prevent backflow of blood
Capillary Function Blood flows through only 5 10% of the body s capillaries at any given time Capillaries in major organs are usually filled to capacity Blood supply varies in many other sites Two mechanisms regulate distribution of blood in capillary beds Constriction or dilation of arterioles that supply capillary beds Precapillary sphincters that control flow of blood between arterioles and venules Blood flow is regulated by nerve impulses, hormones, and other chemicals
Figure 42.13 Precapillary sphincters Thoroughfare channel Arteriole (a) Sphincters relaxed Capillaries Venule Arteriole (b) Sphincters contracted Venule
The exchange of substances between the blood and interstitial fluid takes place across the thin endothelial walls of the capillaries The difference between blood pressure and osmotic pressure drives fluids out of capillaries at the arteriole end and into capillaries at the venule end Most blood proteins and all blood cells are too large to pass through the endothelium
Figure 42.14 INTERSTITIAL FLUID Net fluid movement out Body cell Blood pressure Osmotic pressure Arterial end of capillary Direction of blood flow Venous end of capillary
Fluid Return by the Lymphatic System The lymphatic system returns fluid that leaks out from the capillary beds Fluid lost by capillaries is called lymph The lymphatic system drains into veins in the neck Valves in lymph vessels prevent the backflow of fluid
Edema is swelling caused by disruptions in the flow of lymph Lymph nodes are organs that filter lymph and play an important role in the body s defense Lymph nodes When the body is fighting an infection, lymph nodes become swollen and tender
4. Components of Blood
Blood components function in exchange, transport, and defense With open circulation, the fluid is continuous with the fluid surrounding all body cells The closed circulatory systems of vertebrates contain a more highly specialized fluid called blood Blood Composition and Function Blood in vertebrates is a connective tissue consisting of several kinds of cells suspended in a liquid matrix called plasma The cellular elements occupy about 45% of the volume of blood
Figure 42.16 Plasma 55% Constituent Major functions Cellular elements 45% Water Ions (blood electrolytes) Sodium Potassium Calcium Magnesium Chloride Bicarbonate Solvent Osmotic balance, ph buffering, and regulation of membrane permeability Separated blood elements Cell type Leukocytes (white blood cells) Basophils Lymphocytes Number per µl (mm 3 ) of blood 5,000 10,000 Functions Defense and immunity Plasma proteins Albumin Osmotic balance, ph buffering Eosinophils Immunoglobulins (antibodies) Apolipoproteins Fibrinogen Defense Lipid transport Clotting Neutrophils Platelets Monocytes 250,000 400,000 Blood clotting Substances transported by blood Nutrients (such as glucose, fatty acids, vitamins) Waste products of metabolism Respiratory gases (O 2 and CO 2 ) Hormones Erythrocytes (red blood cells) 5,000,000 6,000,000 Transport of O 2 and some CO 2
Plasma Plasma contains inorganic salts as dissolved ions, sometimes called electrolytes Plasma proteins influence blood ph and help maintain osmotic balance between blood and interstitial fluid Particular plasma proteins function in lipid transport, immunity, and blood clotting Plasma is similar in composition to interstitial fluid, but plasma has a much higher protein concentration
Cellular Elements Suspended in blood plasma are two types of cells Red blood cells (erythrocytes) transport O 2 White blood cells (leukocytes) function in defense Platelets are fragments of cells that are involved in clotting
Erythrocytes Red blood cells, or erythrocytes, are the most numerous blood cells They contain hemoglobin, the iron-containing protein that transports four molecules of O 2 In mammals, mature erythrocytes lack nuclei and mitochondria
Leukocytes There are 5 major types of white blood cells, or leukocytes: monocytes, neutrophils, basophils, eosinophils, and lymphocytes They function in defense either by phagocytizing bacteria and debris or by mounting immune responses against foreign substances They are found both in and outside of the circulatory system
Stem Cells and the Replacement of Cellular Elements Erythrocytes, leukocytes, and platelets all develop from a common source of stem cells in the red marrow of bones, especially ribs, vertebrae, sternum, and pelvis The hormone erythropoietin (EPO) stimulates erythrocyte production when O 2 delivery is low
Figure 42.17 Stem cells (in bone marrow) Lymphoid stem cells Myeloid stem cells B cells T cells Lymphocytes Erythrocytes Neutrophils Basophils Monocytes Platelets Eosinophils
Figure 42.18 Blood Clotting A cascade of complex reactions converts inactive fibrinogen to fibrin, forming a clot 1 2 3 Collagen fibers Platelet Platelet plug Fibrin clot Red blood cells caught in threads of fibrin 5 µm Clotting factors from: Platelets Damaged cells Plasma (factors include calcium, vitamin K) Fibrin clot formation Enzymatic cascade Prothrombin Thrombin Fibrinogen Fibrin
Cardiovascular Disease Cardiovascular diseases are disorders of the heart and the blood vessels These diseases range in seriousness from minor disturbances of vein or heart function to life-threatening disruptions of blood flow to the heart or brain Atherosclerosis, Heart Attacks, and Stroke One type of cardiovascular disease, atherosclerosis, is caused by the buildup of fatty deposits (plaque) within arteries Cholesterol is a key player in the development of atherosclerosis
Low-density lipoprotein (LDL) delivers cholesterol to cells for membrane production High-density lipoprotein (HDL) scavenges excess cholesterol for return to the liver Risk for heart disease increases with a high LDL to HDL ratio Inflammation is also a factor in cardiovascular disease Endothelium Lumen Thrombus Plaque
A heart attack, or myocardial infarction, is the damage or death of cardiac muscle tissue resulting from blockage of one or more coronary arteries A stroke is the death of nervous tissue in the brain, usually resulting from rupture or blockage of arteries in the head Angina pectoris is chest pain caused by partial blockage of the coronary arteries
Risk Factors and Treatment of Cardiovascular Disease A high LDL/HDL ratio increases the risk of cardiovascular disease The proportion of LDL relative to HDL can be decreased by exercise and by avoiding smoking and foods with trans fats Drugs called statins reduce LDL levels and risk of heart attacks
Inflammation plays a role in atherosclerosis and thrombus formation Aspirin inhibits inflammation and reduces the risk of heart attacks and stroke Hypertension, or high blood pressure, also contributes to heart attack and stroke, as well as other health problems Hypertension can be controlled by dietary changes, exercise, and/or medication
5. Gas Exchange
Gas exchange occurs across specialized respiratory surfaces Gas exchange supplies O 2 for cellular respiration and disposes of CO 2
Partial Pressure Gradients in Gas Exchange Partial pressure is the pressure exerted by a particular gas in a mixture of gases Partial pressures also apply to gases dissolved in liquids such as water Gases undergo net diffusion from a region of higher partial pressure to a region of lower partial pressure
Respiratory Media Animals can use air or water as the O 2 source, or respiratory medium In a given volume, there is less O 2 available in water than in air Obtaining O 2 from water requires greater efficiency than air breathing
Respiratory Surfaces Animals require large, moist respiratory surfaces for exchange of gases between their cells and the respiratory medium, either air or water Gas exchange across respiratory surfaces takes place by diffusion Respiratory surfaces vary by animal and can include the skin, gills, tracheae, and lungs
Ventilation moves the respiratory medium over the respiratory surface Aquatic animals move through water or move water over their gills for ventilation Coelom Parapodium (functions as gill) (a) Marine worm Gills (b) Crayfish Gills Tube foot (c) Sea star
Fish gills use a countercurrent exchange system, where blood flows in the opposite direction to water passing over the gills; blood is always less saturated with O 2 than the water it meets In fish gills, more than 80% of the O 2 dissolved in the water is removed as water passes over the respiratory surface
Figure 42.22 Gill arch O 2 -poor blood O 2 -rich blood Blood vessels Gill arch Water flow Operculum Lamella Water flow Blood flow Countercurrent exchange Gill filaments P O 2 (mm Hg) in water 150 120 90 60 30 Net diffusion of O 2 PO (mm Hg) 140 110 80 50 2 in blood 20
2.5 µm Tracheal Tracheoles Systems Mitochondria Muscle in Insects fiber The tracheal system of insects consists of a network of branching tubes throughout the body The tracheal tubes supply O 2 directly to body cells Air sacs Tracheae Air sac Tracheole Body cell Trachea External opening Air
Lungs Lungs are an infolding of the body surface The circulatory system (open or closed) transports gases between the lungs and the rest of the body The size and complexity of lungs correlate with an animal s metabolic rate
Figure 42.24 Mammalian Respiratory Systems Pharynx Larynx (Esophagus) Trachea Right lung Bronchus Bronchiole Branch of pulmonary vein (oxygen-rich blood) Terminal bronchiole Nasal cavity Left lung Capillaries Branch of pulmonary artery (oxygen-poor blood) Alveoli 50 µm Diaphragm (Heart) Dense capillary bed enveloping alveoli (SEM)
Air inhaled through the nostrils is filtered, warmed, humidified, and sampled for odors The pharynx directs air to the lungs and food to the stomach Swallowing moves the larynx upward and tips the epiglottis over the glottis in the pharynx to prevent food from entering the trachea Cilia and mucus line the epithelium of the air ducts and move particles up to the pharynx This mucus escalator cleans the respiratory system and allows particles to be swallowed into the esophagus
Gas exchange takes place in alveoli, air sacs at the tips of bronchioles Oxygen diffuses through the moist film of the epithelium and into capillaries Carbon dioxide diffuses from the capillaries across the epithelium and into the air space Alveoli lack cilia and are susceptible to contamination Secretions called surfactants coat the surface of the alveoli Preterm babies lack surfactant and are vulnerable to respiratory distress syndrome; treatment is provided by artificial surfactants
Breathing ventilates the lungs The process that ventilates the lungs is breathing, the alternate inhalation and exhalation of air How an Amphibian Breathes An amphibian such as a frog ventilates its lungs by positive pressure breathing, which forces air down the trachea
How a Bird Breathes Birds have eight or nine air sacs that function as bellows that keep air flowing through the lungs Air passes through the lungs in one direction only Passage of air through the entire system of lungs and air sacs requires two cycles of inhalation and exhalation Ventilation in birds is highly efficient
Figure 42.26 Anterior air sacs Posterior air sacs Lungs Airflow 1 mm Air tubes (parabronchi) in lung Posterior air sacs Lungs 3 Anterior air sacs 2 4 1 1 2 First inhalation First exhalation 3 4 Second inhalation Second exhalation
How a Mammal Breathes Mammals ventilate their lungs by negative pressure breathing, which pulls air into the lungs Lung volume increases as the rib muscles and diaphragm contract The tidal volume is the volume of air inhaled with each breath The maximum tidal volume is the vital capacity After exhalation, a residual volume of air remains in the lungs
Figure 42.27 Rib cage expands as rib muscles contract. Rib cage gets smaller as rib muscles relax. Lung Diaphragm 1 INHALATION: Diaphragm contracts (moves down). 2 EXHALATION: Diaphragm relaxes (moves up).
Control of Breathing in Humans In humans, breathing is usually regulated by involuntary mechanisms The breathing control centers are found in the medulla oblongata of the brain The medulla regulates the rate and depth of breathing in response to ph changes in the cerebrospinal fluid
Figure 42.28 Blood CO 2 level falls and ph rises. NORMAL BLOOD ph (about 7.4) Medulla detects decrease in ph of cerebrospinal fluid. Blood ph falls due to rising levels of CO 2 in tissues (such as when exercising). Signals from medulla to rib muscles and diaphragm increase rate and depth of ventilation. Cerebrospinal fluid Medulla oblongata Carotid arteries Aorta Medulla receives signals from major blood vessels. Sensors in major blood vessels detect decrease in blood ph.
Sensors in the aorta and carotid arteries monitor O 2 and CO 2 concentrations in the blood These signal the breathing control centers, which respond as needed Additional modulation of breathing takes place in the pons, next to the medulla
Coordination of Circulation and Gas Exchange Blood arriving in the lungs has a low partial pressure of O 2 and a high partial pressure of CO 2 relative to air in the alveoli In the alveoli, O 2 diffuses into the blood and CO 2 diffuses into the air In tissue capillaries, partial pressure gradients favor diffusion of O 2 into the interstitial fluids and CO 2 into the blood
Figure 42.29 120 160 27 6 Exhaled air Inhaled air 1 0.2 P O2 P CO2 P O2 P CO2 40 45 5 Alveolar epithelial cells Blood entering alveolar capillaries CO 2 O 2 Alveolar spaces Alveolar capillaries 2 104 P O2 40 P CO2 P O2 P CO2 Pulmonary arteries and systemic veins Pulmonary veins and systemic arteries 3 104 40 P O2 P CO2 <40 >45 P O2 P CO2 4 Body tissue CO 2 O 2 Systemic capillaries
Respiratory Pigments Respiratory pigments, proteins that transport oxygen, greatly increase the amount of oxygen that blood can carry Arthropods and many molluscs have hemocyanin with copper as the oxygen-binding component Most vertebrates and some invertebrates use hemoglobin In vertebrates, hemoglobin is contained within erythrocytes
A single hemoglobin molecule can carry four molecules of O 2, one molecule for each ironcontaining heme group The hemoglobin dissociation curve shows that a small change in the partial pressure of oxygen can result in a large change in delivery of O 2 Hemoglobin Iron Heme
O 2 saturation of hemoglobin (%) O 2 saturation of hemoglobin (%) Figure 42.30 100 80 60 40 O 2 unloaded to tissues at rest O 2 unloaded to tissues during exercise 100 80 60 40 ph 7.4 ph 7.2 Hemoglobin retains less O 2 at lower ph (higher CO 2 concentration) 20 20 0 0 20 Tissues during exercise 40 60 Tissues at rest (mm Hg) P O 2 80 100 Lungs (a) P O and hemoglobin dissociation at ph 7.4 2 0 0 20 40 60 80 P O 2 (mm Hg) (b) ph and hemoglobin dissociation 100 CO 2 produced during cellular respiration lowers blood ph and decreases the affinity of hemoglobin for O 2 ; this is called the Bohr shift Hemoglobin plays a minor role in transport of CO 2 and assists in buffering the blood
Carbon Dioxide Transport Some CO 2 from respiring cells diffuses into the blood and is transported in blood plasma, bound to hemoglobin The remainder diffuses into erythrocytes and reacts with water to form H 2 CO 3, which dissociates into H + and bicarbonate ions (HCO 3 ) In the lungs the relative partial pressures of CO 2 favor the net diffusion of CO 2 out of the blood
Respiratory Adaptations of Diving Mammals Diving mammals have evolutionary adaptations that allow them to perform extraordinary feats For example, Weddell seals in Antarctica can remain underwater for 20 minutes to an hour For example, elephant seals can dive to 1,500 m and remain underwater for 2 hours These animals have a high blood to body volume ratio Weddell seal
Deep-diving air breathers stockpile O 2 and deplete it slowly Diving mammals can store oxygen in their muscles in myoglobin proteins Diving mammals also conserve oxygen by Changing their buoyancy to glide passively Decreasing blood supply to muscles Deriving ATP in muscles from fermentation once oxygen is depleted