Circulation and Gas Exchange

Open and Closed Circulatory Systems Both systems have three basic components: A circulatory fluid (blood or hemolymph) A set of tubes (blood vessels) A muscular pump (the heart) open circulatory system In insects, other arthropods, and most molluscs blood bathes the organs directly no distinction between blood and interstitial fluid (hemolymph)

LE 42-3 Heart Heart Hemolymph in sinuses surrounding organs Interstitial fluid Small branch vessels in each organ Anterior vessel Lateral vessel Ostia Dorsal vessel (main heart) Tubular heart Auxiliary hearts Ventral vessels An open circulatory system. A closed circulatory system.

closed circulatory system blood is confined to vessels and is distinct from the interstitial fluid

Arteries carry blood to capillaries where chemical exchange between the blood and interstitial fluid Veins return blood from capillaries to the heart

Fishes 2 chambered heart Gills for gas exchange one ventricle and one atrium Gills for gas exchange

Amphibians 3 chambered heart two atria and one ventricle

Reptiles (Except Birds) double circulation pulmonary circuit (lungs) systemic circuit 3 chambered heart

Mammals and Birds 4 chambered heart 2 atria and 2 ventricle left side receives oxygen-rich blood right side receives oxygen-poor blood

A powerful four-chambered heart was an essential adaptation of the endothermic way of life characteristic of mammals and birds

REPTILES (EXCEPT BIRDS) Lung and skin capillaries FISHES AMPHIBIANS REPTILES (EXCEPT BIRDS) MAMMALS AND BIRDS Gill capillaries Lung and skin capillaries Lung capillaries Lung capillaries Gill circulation Pulmocutaneous circuit Right systemic aorta Pulmonary circuit Pulmonary circuit Artery Heart: Ventricle (V) Left systemic aorta A A A A A A Atrium (A) V V V V V Right Left Right Left Right Left Systemic circulation Systemic circuit Systemic circuit Vein Systemic capillaries Systemic capillaries Systemic capillaries Systemic capillaries Systemic circuits include all body tissues except lungs. Note that circulatory systems are depicted as if the animal is facing you: with the right side of the heart shown at the left and vice-versa.

LE 42-5 Anterior vena cava Capillaries of head and forelimbs Pulmonary artery Pulmonary artery Capillaries of right lung Aorta Capillaries of left lung Pulmonary vein Pulmonary vein Right atrium Left atrium Right ventricle Left ventricle Posterior vena cava Aorta Capillaries of abdominal organs and hind limbs

Pulmonary artery Aorta Anterior vena cava Pulmonary artery Right LE 42-6 Pulmonary artery Aorta Anterior vena cava Pulmonary artery Right atrium Left atrium Pulmonary veins Pulmonary veins Semilunar valve Semilunar valve Atrioventricular valve Atrioventricular valve Posterior vena cava 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

LE 42-7 Atrial systole; ventricular Semilunar diastole valves closed 0.1 sec Semilunar valves open AV valves open 0.3 sec 0.4 sec Atrial and ventricular diastole AV valves closed Ventricular systole; atrial diastole

The heart rate, also called the pulse, is the number of beats per minute The cardiac output is the volume of blood pumped into the systemic circulation per minute

Maintaining the Heart’s Rhythmic Beat Some cardiac muscle cells are self-excitable, meaning they contract without any signal from the nervous system

The sinoatrial (SA) node, or pacemaker, sets the rate and timing at which cardiac muscle cells contract 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

Impulses that travel during the cardiac cycle can be recorded as an electrocardiogram (ECG or EKG)

LE 42-8 Pacemaker generates wave of signals to contract. Signals are delayed at AV node. Signals pass to heart apex. Signals spread throughout ventricles. SA node (pacemaker) AV node Bundle branches Purkinje fibers Heart apex ECG

The pacemaker is influenced by nerves, hormones, body temperature, and exercise

Concept 42.3: Physical principles govern blood circulation The physical principles that govern movement of water in plumbing systems also influence the functioning of animal circulatory systems

Blood Vessel Structure and Function The “infrastructure” of the circulatory system is its network of blood vessels All blood vessels are built of similar tissues and have three similar layers

LE 42-9 Artery Vein 100 µm Endothelium Valve Basement membrane Smooth muscle Smooth muscle Capillary Connective tissue Connective tissue Artery Vein Arteriole Venule

Structural differences in arteries, veins, and capillaries correlate with functions Arteries have thicker walls that 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

LE 42-10 Direction of blood flow in vein (toward heart) Valve (open) Skeletal muscle Valve (closed)

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

LE 42-11 5,000 4,000 Area (cm2) 3,000 2,000 1,000 50 40 Velocity (cm/sec) 30 20 10 120 Systolic pressure 100 80 Pressure (mm Hg) 60 Diastolic pressure 40 20 Aorta Arteries Arterioles Capillaries Venules Veins Venae cavae

Blood Pressure Blood pressure is the hydrostatic pressure that blood exerts against the wall of a vessel 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 Blood pressure is determined by cardiac output and peripheral resistance due to constriction of arterioles

LE 42-12_4 Blood pressure reading: 120/70 Pressure in cuff above 120 below 120 Pressure in cuff below 70 Rubber cuff inflated with air 120 120 70 Sounds audible in stethoscope Sounds stop Artery Artery closed

Capillary Function 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: Contraction of the smooth muscle layer in the wall of an arteriole constricts the vessel Precapillary sphincters control flow of blood between arterioles and venules

LE 42-13ab Precapillary sphincters Thoroughfare channel Arteriole Venule Capillaries Sphincters relaxed Arteriole Venule Sphincters contracted

Capillaries and larger vessels (SEM) 20 µm LE 42-13c Capillaries and larger vessels (SEM) 20 µm

The critical 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

LE 42-14 Tissue cell INTERSTITIAL FLUID Net fluid movement out Net fluid movement in Capillary Capillary Red blood cell 15 µm Direction of blood flow Blood pressure Osmotic pressure Inward flow Pressure Outward flow Arterial end of capillary Venous end

Fluid Return by the Lymphatic System The lymphatic system returns fluid to the body from the capillary beds This system aids in body defense Fluid reenters the circulation directly at the venous end of the capillary bed and indirectly through the lymphatic system

Concept 42.4: Blood is a connective tissue with cells suspended in plasma In invertebrates with open circulation, blood (hemolymph) is not different from interstitial fluid Blood in the circulatory systems of vertebrates is a specialized connective tissue

Blood Composition and Function Plasma Fluid 55% of blood composition Cellular Components 45% of blood composition

Plasma Blood plasma is about 90% water Solutes inorganic salts dissolved ions sometimes called electrolytes plasma proteins influence blood pH, osmotic pressure, and viscosity lipid transport Immunity blood clotting Ex. albumins, globulins, fibrinogen

LE 42-15 Plasma 55% Constituent Major functions Cellular elements 45% Water Solvent for carrying other substances Cell type Number Functions per µL (mm3) of blood Erythrocytes (red blood cells) Ions (blood electrolytes) 5–6 million Transport oxygen and help transport carbon dioxide Sodium Potassium Calcium Magnesium Chloride Bicarbonate Osmotic balance, pH buffering, and regulation of membrane permeability Separated blood elements Leukocytes (white blood cells) Defense and immunity 5,000–10,000 Plasma proteins Albumin Osmotic balance, pH buffering Lymphocyte Fibrinogen Clotting Basophil Immunoglobulins (antibodies) Defense Eosinophil Substances transported by blood Neutrophil Monocyte Nutrients (such as glucose, fatty acids, vitamins) Waste products of metabolism Respiratory gases (O2 and CO2) Hormones Platelets 250,000– 400,000 Blood clotting

Cellular Elements Suspended in blood plasma are two types of cells: Red blood cells (erythrocytes) transport oxygen Most abundant White blood cells (leukocytes) function in defense Include monocytes, neutrophils, basophils, eosinophils, and lymphocytes defense phagocytizing bacteria and debris producing antibodies Platelets involved in clotting

Erythrocytes, leukocytes, and platelets all develop from a common source, pluripotent stem cells in the red marrow of bones

LE 42-16 Pluripotent stem cells (in bone marrow) Lymphoid stem cells Myeloid stem cells Basophils B cells T cells Lymphocytes Eosinophils Neutrophils Erythrocytes Platelets Monocytes

Blood Clotting When the endothelium of a blood vessel is damaged, the clotting mechanism begins A cascade of complex reactions converts fibrinogen to fibrin, forming a clot

LE 42-17 Endothelium of vessel is damaged, exposing connective tissue; platelets adhere Platelets form a plug Seal is reinforced by a clot of fibrin Collagen fibers Platelet plug Fibrin clot Red blood cell Platelet releases chemicals that make nearby platelets sticky Clotting factors from: Platelets Damaged cells Plasma (factors include calcium, vitamin K) Prothrombin Thrombin Fibrinogen Fibrin 5 µm

One type of cardiovascular disease, atherosclerosis, is caused by the buildup of cholesterol within arteries

Connective tissue Smooth muscle Endothelium Plaque Normal artery 50 µm Partly clogged artery 250 µm

Hypertension, or high blood pressure, promotes atherosclerosis and increases the risk of heart attack and stroke A heart attack is the 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

Concept 42.5: Gas exchange occurs across specialized respiratory surfaces Gas exchange supplies oxygen for cellular respiration and disposes of carbon dioxide Animals require large, moist respiratory surfaces for adequate diffusion of gases between their cells and the respiratory medium, either air or water

Energy-rich fuel molecules from food Respiratory medium (air or water) O2 CO2 Respiratory surface Organismal level Circulatory system Cellular level Energy-rich fuel molecules from food Cellular respiration ATP

Gills in Aquatic Animals Gills are outfoldings of the body surface specialized for gas exchange

In some invertebrates, gills have a simple shape and are distributed over much of the body

LE 42-20a Gills Coelom Tube foot Sea star

Many segmented worms have flaplike gills that extend from each segment of their body

LE 42-20b Parapodia Gill Marine worm

The gills of clams, crayfish, and many other animals are restricted to a local body region

LE 42-20c Gills Scallop

LE 42-20d Gills Crayfish

Effectiveness of gas exchange in some gills, including those of fishes, is increased by ventilation and the countercurrent flow of blood and water

LE 42-21 Oxygen-poor blood Lamella Oxygen-rich blood Gill arch Gill vessel 15% 40% Water flow 70% 5% 30% Operculum 100% 60% 90% Water flow over lamellae showing % O2 O2 Blood flow through capillaries in lamellae showing % O2 Gill filaments Countercurrent exchange

Tracheal Systems in Insects The tracheal system of insects consists of tiny branching tubes that penetrate the body

LE 42-22a Tracheae Air sacs Spiracle

Body cell Air sac Tracheole Trachea Air Body wall Tracheoles LE 42-22b Body cell Air sac Tracheole Trachea Air Body wall Tracheoles Mitochondria Myofibrils 2.5 µm

The tracheal tubes supply O2 directly to body cells

Lungs Spiders, land snails, and most terrestrial vertebrates have internal lungs

Mammalian Respiratory Systems: A Closer Look A system of branching ducts conveys air to the lungs Air inhaled through the nostrils passes through the pharynx into the trachea, bronchi, bronchioles, and dead-end alveoli, where gas exchange occurs

LE 42-23 Branch from Branch pulmonary from artery pulmonary (oxygen-poor blood) Branch from pulmonary vein (oxygen-rich blood) Terminal bronchiole Nasal cavity Pharynx Alveoli Larynx Left lung Esophagus 50 µm Trachea Right lung 50 µm Bronchus Bronchiole Diaphragm Heart SEM Colorized SEM

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 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

LE 42-24 Rib cage gets smaller as rib muscles relax Rib cage expands as rib muscles contract Air inhaled Air exhaled Lung Diaphragm INHALATION Diaphragm contracts (moves down) EXHALATION Diaphragm relaxes (moves up)

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 Every exhalation completely renews the air in the lungs

Air sacs empty; lungs fill LE 42-25 Air Air Anterior air sacs Trachea Posterior air sacs Lungs Lungs Air tubes (parabronchi) in lung 1 mm INHALATION Air sacs fill EXHALATION Air sacs empty; lungs fill

Control of Breathing in Humans In humans, the main breathing control centers are in two regions of the brain, the medulla oblongata and the pons The medulla regulates the rate and depth of breathing in response to pH changes in the cerebrospinal fluid The medulla adjusts breathing rate and depth to match metabolic demands

Sensors in the aorta and carotid arteries monitor O2 and CO2 concentrations in the blood These sensors exert secondary control over breathing

LE 42-26 Cerebrospinal fluid Pons Breathing control centers Medulla oblongata Carotid arteries Aorta Diaphragm Rib muscles

Concept 42.7: Respiratory pigments bind and transport gases The metabolic demands of many organisms require that the blood transport large quantities of O2 and CO2

The Role of Partial Pressure Gradients Gases diffuse down pressure gradients in the lungs and other organs Diffusion of a gas depends on differences in a quantity called partial pressure

A gas diffuses from a region of higher partial pressure to a region of lower partial pressure In the lungs and tissues, O2 and CO2 diffuse from where their partial pressures are higher to where they are lower

LE 42-27 Inhaled air Exhaled air Alveolar spaces O2 CO2 O2 CO2 160 0.2 120 27 Alveolar spaces O2 CO2 O2 CO2 104 40 Alveolar epithelial cells O2 CO2 CO2 O2 Blood entering alveolar capillaries Blood leaving alveolar capillaries CO2 O2 Alveolar capillaries of lung 40 45 104 40 O2 CO2 O2 CO2 Pulmonary arteries Pulmonary veins Systemic veins Systemic arteries Heart Tissue capillaries CO2 O2 Blood leaving tissue capillaries Blood entering tissue capillaries 40 45 CO2 O2 100 40 O2 CO2 O2 CO2 Tissue cells < 40 > 45 O2 CO2

Respiratory Pigments Respiratory pigments, proteins that transport oxygen, greatly increase the amount of oxygen that blood can carry

Oxygen Transport The respiratory pigment of almost all vertebrates is the protein hemoglobin, contained in erythrocytes Like all respiratory pigments, hemoglobin must reversibly bind O2, loading O2 in the lungs and unloading it in other parts of the body

Heme group Iron atom O2 loaded in lungs O2 unloaded in tissues LE 42-28 Heme group Iron atom O2 loaded in lungs O2 unloaded in tissues Polypeptide chain

Loading and unloading of O2 depend on cooperation between the subunits of the hemoglobin molecule The binding of O2 to one subunit induces the other subunits to bind O2 with more affinity Cooperative O2 binding and release is evident in the dissociation curve for hemoglobin A drop in pH lowers affinity of hemoglobin for O2

O2 saturation of hemoglobin (%) LE 42-29a 100 O2 unloaded from hemoglobin during normal metabolism 80 O2 saturation of hemoglobin (%) 60 O2 reserve that can be unloaded from hemoglobin to tissues with high metabolism 40 20 20 40 60 80 100 Tissues during exercise Tissues at rest Lungs P (mm Hg) O2 P and hemoglobin dissociation at 37°C and pH 7.4 O2

O2 saturation of hemoglobin (%) LE 42-29b 100 pH 7.4 80 Bohr shift: additional O2 released from hemoglobin at lower pH (higher CO2 concentration) O2 saturation of hemoglobin (%) 60 pH 7.2 40 20 20 40 60 80 100 P (mm Hg) O2 pH and hemoglobin dissociation

Carbon Dioxide Transport Hemoglobin also helps transport CO2 and assists in buffering Carbon from respiring cells diffuses into the blood plasma and then into erythrocytes and is ultimately released in the lungs

LE 42-30 Tissue cell CO2 transport from tissues CO2 produced Interstitial fluid CO2 Blood plasma within capillary CO2 Capillary wall CO2 H2O Red blood cell H2CO3 Carbonic acid Hemoglobin picks up CO2 and H+ Hb HCO3– Bicarbonate + H+ HCO3– To lungs CO2 transport to lungs HCO3– HCO3– + H+ Hemoglobin releases CO2 and H+ H2CO3 Hb H2O CO2 CO2 CO2 CO2 Alveolar space in lung

Animation: O2 From Blood to Tissues Animation: O2 From Tissues to Blood Animation: O2 From Blood to Lungs Animation: O2 From Lungs to Blood

Elite Animal Athletes Migratory and diving mammals have evolutionary adaptations that allow them to perform extraordinary feats

The Ultimate Endurance Runner The extreme O2 consumption of the antelope-like pronghorn underlies its ability to run at high speed over long distances

Diving Mammals Deep-diving air breathers stockpile O2 and deplete it slowly