Chapter Objectives List five cardiovascular system functions Describe interactions among cardiac output, total peripheral resistance, and arterial blood pressure (BP) Chapter 15 The Cardiovascular System Describe how the venous system acts as an active blood reservoir Describe how to measure BP, and list typical systolic and diastolic blood pressure values during rest and physical activity Discuss how BP responds to resistance exercises Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Chapter Objectives, cont. Cardiovascular System Components Explain the hypotensive response during recovery from physical activity • Consists of four components Diagram the major vessels of the coronary circulation Describe the pattern of myocardial blood • flow, VO2, and substrate use during rest and various intensities of physical exertion 1. A pump that provides continuous linkage with the other three components 2. A high-pressure distribution circuit 3. Exchange vessels 4. A low-pressure collection and return circuit Explain the rate–pressure product, its meaning, and rationale for use in clinical exercise physiology Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Schematic View of the Cardiovascular System Showing Heart and Pulmonary and Systemic Vascular Circuits Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins The Heart • Four-chambered organ that provides the drive for blood flow • Weighs 11 oz. for average male, 9 oz. for female • Pumps ~70mL for each beat (stroke volume) at rest At rest in one day, ~1900 gallons pumped through the heart, or 52 million gallons for a 75-y life span • Heart muscle called myocardium; myocardial fibers interconnect in latticework fashion to allow the heart to function as a unit Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 1 The Heart, cont. The Heart’s Valves • Right side 1. Atrioventricular valves: Receive blood returning from body Pumps blood to lungs for aeration through pulmonary circulation • Left side Receives oxygenated blood from lungs Pumps blood into thick-walled muscular aorta for distribution via systemic circulation Tricuspid: provides one-way blood flow from the right atrium to right ventricle Bicuspid/Mitral: provides one-way blood flow from left atrium to left ventricle 2. Semilunar valves: Located in arterial wall just outside heart; prevent blood from flowing back into the heart between contractions • Two sides separated by interventricular septum Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Myocardial Contraction • Atrial chambers serve as “primer pumps” to receive and store blood during ventricular contraction • About 70% of blood returning to atria flows directly into ventricles before atrial contraction • Simultaneous contraction of both atria forces remaining blood into ventricles • Almost immediately after atrial contraction, ventricles contract and propel blood into arterial system Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins The Heart, its Vessels, and Flow of Blood High-pressure tubing that propels O2-rich blood to tissues Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins The Arterial System • High-pressure tubing that propels oxygen-rich blood to tissues • Comprised of layers of connective tissue and smooth muscle • No gaseous exchange occurs between arterial blood and surrounding tissues • Blood pumped from left ventricle enters aorta and is distributed throughout the body through a network of arteries and arterioles Smooth muscle in arteriole walls either constrict or relax to regulate blood flow to periphery Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Blood Pressure • Force of blood against arterial walls during cardiac cycle • Peripheral vessels do not permit blood to “run off” into arterial system as rapidly as it ejects from heart • Aorta “stores” some ejected blood, which creates pressure within the entire arterial system • Arterial blood pressure reflects the combined effects of arterial blood flow per minute and resistance to flow in peripheral vasculature Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 2 Blood Pressure, cont. Blood Pressure, cont. • Systolic Blood Pressure (SBP) Mean Arterial Pressure (MAP) = DBP + [0.333 (SBP – DBP)] Provides estimate of work of heart and force blood exerts against arterial walls during systole • Diastolic Blood Pressure (DBP): relaxation phase of cardiac cycle (diastole) Indicates peripheral resistance or ease that blood flows from arterioles into capillaries • Mean Arterial Pressure (MAP): average force exerted by blood against arterial wall during cardiac cycle Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Adult Blood Pressure Classification Values Classification Normal Systolic BP (mm Hg) Diastolic BP (mm Hg) <120 and <80 Prehypertension 120-139 or 80-89 Stage 1 Hypertension 140-159 or 90-99 Stage 2 Hypertension ≥160 or ≥100 Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Structure of Blood Vessels Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Cardiac Output (Q) and Total Peripheral Resistance (TPR) Q = MAP ÷ TPR MAP and cardiac output estimate change in total resistance to blood flow in the transition from rest to exercise TPR = MAP ÷ Q Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Blood Flow in Capillaries • The precapillary sphincter consists of a ring of smooth muscle that encircles the capillary at its origin and controls its diameter • Sphincter constriction and relaxation provide a means for blood flow regulation within a specific tissue to meet metabolic requirements • Two factors trigger precapillary sphincter relaxation to open more capillaries: 1. Driving force of increased local BP plus intrinsic neural control 2. Local metabolites produced in exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 3 Capillary Blood Flow During Rest [A] and Exercise [B] The Venous System • Capillaries feed deoxygenated blood into small veins or venules • Veins in lower body eventually empty into inferior vena cava, the body’s largest vein Vena cava returns blood to right atrium from abdomen, pelvis, and lower extremities • Venous blood from vessels in head, neck, shoulder region, thorax, and abdominal wall flows into superior vena cava to join inferior vena cava at the heart (mixed-venous blood) Mixed-venous blood enters right atrium Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Venous Return • Valves within veins allow blood to flow in only one direction toward the heart • Smallest muscular contractions or minor pressure changes within the thoracic cavity with breathing readily compressing veins • Alternate compression and relaxation of veins and one-way valve action provide a “milking” action to propel blood back to the heart Without valves, blood would stagnate in extremity veins; people would faint every time they stood up because of reduced venous return and diminished cerebral blood flow Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Blood Flow and BP in Systemic Circulation at Rest • Blood pressure within each portion of the arterial system relates to total area (resistance) in that section Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Valves in Veins • Valves (A) prevent backflow of blood, but (B) do not hinder normal 1-way flow; (C) blood moves through veins by action of nearby active muscle or, (D) contraction of smooth muscle Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Varicose Veins • Valves within a vein fail to maintain one-way blood flow; blood gathers in vein so they become excessively distended and painful • Usually occurs in surface veins of lower extremities • In severe cases, phlebitis occurs where the venous wall becomes inflamed and deteriorates • People with varicose veins should avoid static, straining-type exercises like resistance training • Exercise does not prevent varicose veins, but regular exercise can minimize discomfort and complications Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 4 Hypertension • Systolic or diastolic pressure that exceeds recommended values Percentage of U.S. Population with Hypertension Chronically strains cardiovascular system; if left untreated, leads to arteriosclerosis, heart disease, stroke, and kidney failure • Artery characteristics in hypertension Excessive resistance to peripheral blood flow because of: - Arterial hardening with fatty material in walls - Neural hyperactivity or kidney malfunction Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Hypertension: Effective Treatment Options Advice Lose excess weight Follow a DASH diet Daily physical activity Limit sodium Limit alcohol Details For every 20 lb reduced Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Pharmacologic Therapies for Hypertension Drop in Systolic Blood Pressure 5 to 20 mm Hg Eat a lower-fat diet rich in vegetables, 8 to 14 mm Hg fruits, and low-fat dairy foods Do 30 min a day of aerobic activity like 4 to 9 mm Hg brisk walking Eat no more than 2400 mg a day 2 to 8 mm Hg (1500 mg is better) Consume no more than two drinks 2 to 8 mm Hg daily (men) and one drink daily (women) (one drink = 12 oz. beer, 5 oz. wine, or 1.5 oz. 80-proof whiskey) 2 to 4 mm Hg Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins BP Response to Resistance Exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Blood Pressure During Resistance Exercise • Straining exercises mechanically compress peripheral arteries that supply active muscle Arterial vascular compression dramatically increases total peripheral resistance and reduces muscle perfusion In an attempt to restore muscle blood flow, substantial increases occur in sympathetic nervous system activity, cardiac output, and MAP • Acute cardiovascular strain with heavy resistance exercise could prove harmful to individuals with heart and vascular disease Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 5 BP: Steady-Rate Physical Activity • During rhythmic muscle activity, vasodilation BP: Graded Exercise in active muscles reduces TPR to enhance blood flow through peripheral vasculature • After an initial rapid rise from resting level, • Alternate muscle contraction/relaxation propels • DPB remains stable or decreases slightly at • Increased blood flow during steady-rate exercise • SBP may increase to 200 mm Hg or higher in blood through the vascular circuit back to the heart rapidly increases SBP during the first few minutes • SBP often declines as steady-rate exercise continues because arterioles in active muscles continue to dilate, further reducing peripheral resistance to blood flow; DBP generally remains unchanged throughout exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins SBP and DBP During Graded Exercise SBP increases linearly with exercise intensity higher exercise levels healthy, fit individuals during maximum exercise (despite reduced TPR) This blood pressure most likely reflects the heart’s large cardiac output during maximal exercise by individuals with high aerobic capacity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins BP: Upper-Body Physical Activity • Exercise with arms produces higher SBP and DBP than leg exercise performed at a given • percentage of VO2max in each exercise mode Occurs because smaller arm muscle mass and vasculature offer greater resistance to blood flow than activation of larger leg mass and blood supply • Individuals with cardiovascular dysfunction should rhythmically exercise relatively large muscle groups in contrast to exercise that engages a limited muscle mass Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Recovery From Physical Activity The Heart’s Blood Supply • On completion of a single bout of submaximal • The heart has its own circulatory network, the exercise, BP temporarily falls below pre-exercise levels for normotensive and hypertensive individuals due likely from unexplained peripheral vasodilation • The hypotensive response to exercise can last up to 12 hr Occurs in response to either low- and moderate-intensity aerobic exercise or resistance exercise coronary circulation, that arises from the heart’s top portion • Right and left coronary arteries Greatest blood volume flows in left coronary artery, to left atrium and left ventricle, and right ventricle • Blood leaves tissues of left ventricle through coronary sinus and right ventricle from the anterior cardiac veins • Normal blood flow to myocardium at rest = 200 to 250 mL/min Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 6 Anterior and Posterior View of Coronary Circulation Myocardial Oxygen Supply and Use • At rest, myocardium extracts 70 to 80% of O2 from blood in the coronary vessels • Proportionate increase in coronary blood flow in exercise provides sole mechanism to increase myocardial O2 supply • Two factors increase myocardial blood flow: Elevated myocardial metabolism dilates coronary vessels Increased aortic pressure during exercise forces a proportionately greater volume of blood into coronary circulation Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Impaired Myocardial Blood Supply • Myocardium depends on adequate O2 supply because it has limited anaerobic capacity • Extensive vascular perfusion supplies at least one capillary to each of the heart’s muscle fibers • Tissue hypoxia provides a potent stimulus to myocardial blood flow Can produce chest pains, or angina pectoris Plaque or Blood Clot • Plaque (A) or blood clot (thrombus) (B) lodged in a coronary vessel impairs normal heart function and can result in a myocardial infarction • Exercise provides an effective way to evaluate adequacy of myocardial blood flow A blood clot in a coronary vessel can impair normal heart function, leading to a myocardial infarction Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Myocardial Metabolism Rate-Pressure Product (RPP) • • Estimate of myocardial workload and VO2 • Computed from product of peak SBP measured at the brachial artery and heart rate (HR) Index of relative cardiac work - relates closely to directly measured myocardial VO2 and coronary blood flow in healthy subjects over a wide range of exercise intensities RPP = SBP x HR Ranges from 6000 at rest to ≥40,000 during exercise, depending on intensity and mode Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins • Relies almost exclusively on aerobic energy release • Contains greatest mitochondrial concentration of all tissues • Glucose, fatty acids, and lactate from glycolysis in skeletal muscle provide myocardial energy At rest, myocardial energy comes mainly from FFA Following a meal, the myocardium “prefers” glucose as an energy substrate During intense exercise, myocardium derives its major energy by oxidizing circulating lactate Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 7 Generalized Pattern of Myocardial Substrate Use Summary 1. Myocardium is composed of striated muscle fibers 2. Heart functions as two separate pumps: one pump receives blood from the body and pumps it to the lungs for aeration (pulmonary circulation); the other receives oxygenated blood from the lungs and pumps it throughout systemic circulation 3. Pressure changes created during the cardiac cycle act on the heart’s valves to provide one-way blood flow in the vascular circuit 4. Surge of blood with ventricular contraction and subsequent runoff of blood in relaxation creates pressure changes within the arterial vessels Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Summary, cont. Summary, cont. 5. Ventricular contraction generates systolic SBP 9. At rest, the myocardium extracts approximately • SBP increases in proportion to VO2 and blood flow during graded exercise; DBP remains relatively unchanged or decreases slightly 6. DBP represents lowest BP in relaxation before the next systole 7. Compression and relaxation of veins by muscle action impart energy to facilitate venous return. This “muscle pump” mechanism provides justification for active recovery immediately following vigorous effort 8. Abnormal elevated BP (hypertension) imposes a chronic cardiovascular stress that damages arterial vessels and leads to arteriosclerosis, heart disease, stroke, and kidney failure Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 10.At same relative and absolute exercise levels, upper-body exercise produces greater rise in SBP than leg exercise 11.Following physical activity, BP decreases below pre-exercise level and can remain lower for up to 12 h Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Summary, cont. 12.Performing intense resistance exercises poses a risk to individuals with hypertension or other forms of heart disease 13.At rest, the myocardium extracts approximately 80% of O2 flowing through the coronary arteries 14.The rate-pressure product (HR x SBP) estimates myocardial workload 15.Percentage myocardial use of macronutrients for energy varies with physical activity severity and duration and the individual’s training status Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Chapter 16 Cardiovascular Regulation and Integration Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 8 Chapter Objectives Chapter Objectives, cont. Explain intrinsic and extrinsic factors that regulate heart rate (HR) during rest and physical activity Describe the effects of aerobic training on neural regulation of heart rate Draw and identify the major components of a normal electrocardiogram (ECG) Describe local metabolic factors that regulate blood flow during rest and physical activity Explain “central command” in cardiovascular regulation during exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Distinguish between chemoreceptors, mechanoreceptors, and the metaboreflex in cardiovascular regulation during activity List three factors that affect blood flow Describe two differences in blood flow dynamics to different tissues at exercise onset and during exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Chapter Objectives, cont. Intrinsic Regulation of Heart Rate Describe how nitric oxide regulates local blood flow • Unlike other tissues, cardiac muscle maintains Outline the cardiovascular response to physical activity by a heart transplant patient • Left to its inherent rhythmicity, the heart would its own rhythm beat ~100 b/min • The sinoatrial (SA) node provides the innate stimulus for heat action SA node is referred to as the heart’s pacemaker Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Normal Route for Impulse Transmission Within the Myocardium Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins The Heart’s Electrical Activity • In right atrium the SA node spontaneously depolarizes and repolarizes • The impulse spreads across the atria to atrioventricular (AV) node • A 0.10-s delay occurs to allow atria to contract and propel blood into ventricles Normal route for excitation and conduction of cardiac impulse Time sequence for impulse transmission from SA node though the myocardium Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 9 The Heart’s Electrical Activity, cont. • The AV node gives rise to the AV bundle, also called the bundle of His • The AV bundle transmits the impulse to the Purkinje system that penetrates the right and left ventricles • The transmission of the cardiac impulse flows In the following direction: SA node Atria Purkinje fibers AV node Ventricles Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Normal Transmission of Electrical Impulse Through the Myocardium Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins ECG Waves • P wave: depolarization of atria before atria contract • QRS complex: Signals electrical changes from ventricular depolarization before ventricles contract Atrial repolarization follows P wave, but produces a wave so small that QRS complex usually obscures it • T wave: Represents ventricular repolarization that occurs during ventricular diastole Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Electrocardiogram (ECG) • The myocardium’s electrical activity creates an electrical field throughout the body • The ECG represents a composite record of the heart’s electrical events during a cardiac cycle These electrical events can monitor HR during physical activities and exercise stress testing • A valid ECG tracing requires proper electrode placement ECG leads transmit electrical signal to a recorder, which creates the composite electrical “picture” of myocardial activity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Different Phases of the Normal ECG from Atrial Depolarization to Ventricular Repolarization Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Extrinsic Regulation of Heart Rate and Circulation • Nerves that directly supply the myocardium, and chemical “messengers” that circulate in blood accelerate the heart in anticipation before exercise begins, and then rapidly adjust to intensity of physical effort intensity • Input from the brain and peripheral nervous system continually bombards the cardiovascular control center in the ventrolateral medulla to regulate the heart’s output of blood and blood’s preferential distribution to all body’s tissues Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 10 Neural Mechanisms for Cardiovascular Regulation Before and During Activity Sympathetic and Parasympathetic Neural Input • Neural influences override the myocardium’s inherent rhythm • Originate in cardiovascular center and flow through sympathetic and parasympathetic components of autonomic nervous system Large numbers of sympathetic and parasympathetic neurons innervate the atria, whereas the ventricles receive sympathetic fibers almost exclusively Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Distribution of Sympathetic and Parasympathetic Nerve Fibers Within the Myocardium Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Sympathetic Influence • Stimulation of sympathetic cardioaccelerator nerves releases epinephrine and norepinephrine Cause chronotropic and inotropic effects on heart • Sympathetic stimulation also produces generalized vasoconstriction except in coronary arteries Norepinephrine, released by adrenergic fibers, acts as a vasoconstrictor Dilation of blood vessels under adrenergic influence occurs from decreased adrenergic activity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Chemical, Anatomic, and Functional Organization of Sympathetic and Parasympathetic Divisions of the Autonomic Nervous System Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Parasympathetic Influence • Parasympathetic neurons release acetylcholine, which delays rate of sinus discharge to slow HR • Bradycardia results from stimulation of vagus nerve from medulla’s cardioinhibitory center • Parasympathetic stimulation excites some tissues and inhibits others • At start and during low/moderate intensity exercise, HR increases largely by inhibition of parasympathetic stimulation • HR in strenuous exercise increases by additional parasympathetic inhibition and direct activation of sympathetic cardioaccelerator nerves Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 11 Central Command: Input from Higher Centers • Impulses originating in brain’s higher somatomotor central command center continually modulate medullary activity • Central command provides greatest control over HR during exercise • Heart rapidly “turns on” during exercise by decreasing Influence of Central Command on Heart Rate When Movement Begins parasympathetic inhibitory input and increasing stimulating input from central command • Central command in cardiovascular regulation explains how emotional state can affect cardiovascular response (thus, creating difficulty obtaining “true” resting values for HR and BP) Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Peripheral Input Peripheral Input, cont. • Cardiovascular center receives reflex sensory • Three mechanisms continually assess the input from peripheral receptors in blood vessels, joints, and muscles • Chemoreceptors and mechanoreceptors within muscle and its vasculature monitor its chemical and physical state Modifies either parasympathetic or sympathetic outflow to bring about the appropriate cardiovascular and respiratory responses to various intensities of physical activity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins nature and intensity of exercise and muscle mass activated: 1. Reflex neural input from mechanical deformation of type III afferents within active muscles 2. Chemical stimulation of type IV afferents within active muscles 3. Feed-forward outflow from motor areas of central command Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Peripheral Input, cont. Peripheral Input, cont. • Specific mechanoreceptor feedback governs • Function as negative feedback controllers to: central nervous system’s regulation of blood flow and BP during dynamic exercise Aortic arch and carotid sinus contain pressure-sensitive baroreceptors Inhibit sympathetic outflow from the cardiovascular center Blunt inordinate rise in arterial BP Cardiopulmonary receptors assess mechanical activity in the left ventricle, right atrium, and large veins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 12 Carotid Artery Palpation • External pressure against the carotid artery can slow HR from direct baroreceptor stimulation • Consistently low HR estimation with carotid artery palpation in susceptible individuals would push the person to a higher exercise level. It has no effect in healthy individuals • Vascular disease affects carotid sinus sensitivity and produce low heart rate values Distribution of Blood: Physical Factors Affecting Blood Flow • Blood flow through the vascular circuit follows physical laws of hydrodynamics applied to rigid, cylindrical vessels • Volume of flow in an vessel relates to two factors: 1. Directly to pressure gradient between two ends of the vessels 2. Inversely to resistance encountered to fluid flow • A substitute location measures pulse rate at the radial artery or temporal artery Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Distribution of Blood: Physical Factors Affecting Blood Flow, cont. Distribution of Blood: Physical Factors Affecting Blood Flow, cont. • Friction between blood and internal vascular wall • Poiseuille’s Law: expresses relationship among creates resistance (force) that impedes blood flow • Three factors determine resistance: pressure differential, resistance, and flow Flow = pressure gradient × vessel radius4 ÷vessel length × fluid viscosity 1. Blood thickness (viscosity) The transport vessel length remains constant 2. Length of conducting tube Blood viscosity varies only slightly 3. Blood vessel radius Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins - radius affects blood flow the most - physiologically, constriction and dilation of smaller arterial blood vessels provide mechanism to regulate regional blood flow Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Exercise Effect on Blood Flow Exercise Effect on Blood Flow, cont. • Any increase in energy expenditure requires • Two factors contribute to reduced blood flow rapid adjustments in blood flow that impact the cardiovascular system • During exercise, local arterioles of active muscles dilate while vessels to tissues that can temporarily compromise their blood supply constrict Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins to non-active tissues: 1. Increased sympathetic nervous system outflow 2. Local chemicals that directly stimulate vasoconstriction or enhance effects of other vasoconstrictors Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 13 Factors Within Active Muscle Factors Within Active Muscle, cont. • Skeletal muscle blood flow closely couples to • The opening of dormant capillaries in exercise: metabolic demands Regulation occurs from interaction of neural vasoconstriction activity and locally derived vasoactive substances within endothelium and red blood cells • At rest, 1 in 30 to 40 muscle capillaries Increases total muscle blood flow Delivers a large blood volume with only a minimal increase in blood flow velocity Increases effective surface for gas and nutrient exchange remains open Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Factors Within Active Muscle, cont. • Vasodilation occurs from local factors related to tissue metabolism that act directly on smooth muscle bands of small arterioles and precapillary sphincters Examples include decreased tissue oxygen and local increases in blood flow, temperature, carbon dioxide, acidity, adenosine, magnesium and potassium ions, and nitric oxide production by endothelial cells lining the blood vessels Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Nitric Oxide (NO) and Autoregulation of Tissue Blood Flow • Nitric oxide serves as a signal molecule that dilates blood vessels and decreases vascular resistance • Stimuli from diverse signal chemicals and sheering stress and vessel stretch from increased blood flow through the vessel lumen provoke NO synthesis and release by vascular endothelium In CHD, vascular endothelium produces less NO Venous system may also increase local blood flow by “assessing” increases in the metabolic needs of active muscle and releasing vasodilatory factors Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Nitrous Oxide Regulation of Local Blood Flow Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Integrative Response During Physical Activity • Neural command center above medullary region initiates cardiovascular changes before and during movement • HR and myocardial contractility increase from feed-forward input from cardiovascular center, which depresses parasympathetic activation • Predictable changes in regional blood flow occur in proportion to exercise intensity • Modulation of dilation/constriction optimizes blood flow to needed areas while maintaining BP Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 14 Integrative Response During Physical Activity, cont. • Local metabolic factors act directly to dilate resistance vessels in active muscle • Vasodilation reduces peripheral resistance for greater blood flow in active areas • Vascular constriction in inactive tissues maintains adequate perfusion within active muscle, while maintaining the blood supply to satisfy local metabolic demands of the inactive areas Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Summary, cont. 6. Input from peripheral receptors in blood vessels, joints, and muscles provides feedback to the cardiovascular center with feedback 7. Neural and hormonal extrinsic factors modify the heart’s inherent rhythm 8. HR can reach 200b/min during maximal exercise 9. Carotid artery palpation accurately accesses HR except in certain pathologic conditions 10.Nerves, hormones, and local metabolic factors act on blood vessels’ internal diameter to regulate blood flow Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Summary 1. The cardiovascular system provides for rapid HR regulation and effective blood distribution throughout the body 2. Cardiac rhythm originates at the SA node 3. The ECG records the sequence of the heart’s electrical events during the cardiac cycle 4. Epinephrine and norepinephrine accelerate HR and increase myocardial contractility; acetylcholine acts through the vagus nerve to slow HR 5. The heart turns on in the transition from rest to physical activity from increased sympathetic and decreased parasympathetic activity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Summary, cont. 11.Blood flow changes with the vessels’ radius raised to the fourth power 12.Nitric oxide facilitates blood vessel dilation and decreases vascular resistance 13.Blood flow decreases to kidneys and splanchnic area while muscle blood flow increases during exercise 14.Patients who successfully undergo orthotopic transplantation have a depressed cardiovascular response to exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Chapter Objectives Discuss advantages and disadvantages of methods to measure cardiac output (CO) Compare CO during rest and maximal exercise for the endurance athlete and sedentary person Chapter 17 Functional Capacity of the Cardiovascular System Explain the influence •of each component of the Fick equation on VO2max Discuss physiologic factors that influence SV Contrast components of CO during rest and maximal effort for trained vs. untrained Explain the effects of the Frank-Starling mechanism during physical activity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 15 Chapter Objectives, cont. Cardiac Output Explain cardiovascular drift • Cardiac output expresses the amount of blood Describe cardiac output distribution to major body organs during rest and exercise Describe the relationship between maximum • cardiac output and VO2max along a broad range of fitness levels pumped by the heart during a 1-min period Maximal values reflect the functional capacity of the cardiovascular system Cardiac output = Heart rate × Stroke volume • Methods to assess cardiac output: List three factors that contribute to expanding the a-vO 2 differences during graded exercise Direct Fick Contrast cardiovascular and metabolic dynamics during upper-body versus lower-body exercise CO2 rebreathing Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Indicator dilution Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins The Fick Principle Direct Fick Method • • Expresses relationships between VO2 (mL/min) , - diff (mL/100mL blood) to determine and a-vO 2 cardiac output (mL/min) • - diff Cardiac output = VO ÷ a-vO 2 2 Requires complex invasive methodology • - VO2 (L/min) using open circuit spirometry - average difference between O2 content of arterial and mixed-venous blood (a-vO2diff) Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Indicator Dilution Method CO2 Rebreathing Method • Injects a known quantity of inert dye into • Same open-circuit spirometry method for • a large vein • The indicator material remains in vascular stream and mixes as blood travels to lungs and returns to heart before ejection through systemic circuit • Assessment of arterial blood samples and area under dilution–concentration curve reflects average concentration of blood leaving heart Cardiac Output = Amount dye injected ÷ Average dye concentration in blood for duration of curve × duration of curve Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins determining •VO2 in typical Fick technique but determines VCO2 in the rebreathing method • Using a rapid CO2 gas analyzer and making reasonable assumptions about gas exchange the method provides valid estimates of mixed-venous and arterial CO2 levels Requires breath-by-breath CO2 analysis Does not require blood sampling or medical supervision Can only be used during steady-rate conditions Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 16 Cardiac Output at Rest Cardia Output: Untrained Individuals • Can vary considerably during rest • Resting HR for typical untrained = 70 b/min, • Influencing factors include emotional conditions that alter cortical outflow to cardio-accelerator nerves and nerves that modulate arterial resistance vessels 5 L/min = average cardiac output for males and resting SV = 60 to 70 mL/b • SV and cardiac output for women = 25% below male values Generally relates to a smaller body size - average HR = 70 b/min, average SV = 71 mL/b 4 L/min = average cardiac output for females - average HR = of 70 b/min, average SV = 50 to 60 mL/b Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Cardiac Output: Endurance Athletes • Resting HR of healthy endurance athletes = 50 b/min at rest and resting SV = 100 mL/b • Factors that explain large SV and low HR of endurance-trained athletes: Increased vagal tone and decreased sympathetic drive, both of which slow the heart Increased blood volume, myocardial contractility, and compliance of the left ventricle, all of which augment the heart’s stroke volume Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Cardiac Output During Physical Activity • Systemic blood flow increases directly with intensity of physical activity Cardiac output increases rapidly during the transition from rest to steady-rate exercise and then rises gradually until it plateaus when blood flow meets the exercise metabolic requirements • The endurance athlete achieves a large maximal cardiac output solely through a large SV Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Average Cardiac Output, HR, and SV for Endurance-trained and Untrained Men During Maximal Activity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Maximal Values for Cardiac Output, HR, and SV for Men• with Very Low, Normal, and High VO2max Values Cardiac output = HR x SV • Untrained: 22,000 mL/min = 195 b/min x 113 mL/b • Trained: 35,000 mL/min = 195 b/min x 179 mL/b Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins VO2max (L ∙ min-1) Max Heart Rate (B ∙ min-1) Mitral stenosis 1.6 190 50 9.5 Sedentary 3.2 200 100 20.0 Athlete 5.2 190 160 30.4 Group Max Stroke Volume (mL) Max Cardiac Output (L ∙ min-1) Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 17 Enhancing Stroke Volume: Diastolic Filling Versus Systolic Emptying • Mechanisms that could increase the heart’s SV during exercise: 1. Enhanced cardiac filling in diastole followed by a more forceful systolic contraction 2. Normal ventricular filling with subsequent more forceful ejection and emptying during systole 3. Training adaptations that expand blood volume and reduce resistance to blood flow in peripheral tissues Enhanced Diastolic Filling • Any factor that increases venous return or slows the heart produces greater preload during the cardiac cycle’s diastolic phase An increase in end-diastolic volume stretches myocardial fibers and initiates a more powerful ejection stroke during systole - Ejects the normal SV plus additional blood that entered ventricles during diastole • Frank-Starling law: force of contraction of cardiac muscle remains proportional to its initial resting length Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Greater Systolic Emptying Cardiovascular Drift • Enhanced systolic ejection occurs despite • Describes the gradual time-dependent increased resistance to blood flow in arterial circuit from exercise-induced elevation of systolic BP • Enhanced systolic ejection occurs because ventricles always contain a functional residual blood volume Catecholamine release in exercise enhances myocardial contractile force to augment stroke power and facilitate systolic emptying downward “drift” in cardiovascular responses, most notably SV with concomitant HR increase, during prolonged steady-rate exercise Person must exercise at lower intensity than if cardiovascular drift did not occur • Submaximal exercise for >15 minutes decreases plasma volume, thus decreasing SV Reduced SV initiates a compensatory HR increase to maintain a nearly constant cardiac output Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins SV, HR, and Cerebral Blood Flow (CBF) Under ß-adrenoceptor Blockade and Control Treatments Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Cardiac Output Distribution During Rest • At rest, typical 5-L cardiac output distributes as follows: One fifth flows to muscle tissue (4 to 7 mL/min/100 g muscle) Major portion of remaining blood flows to digestive tract, liver, spleen, brain, and kidneys receive major portions of the remaining blood Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 18 Cardiac Output Distribution During Exercise • Cardiac output diverts to active muscles at maximal exertion Distribution of Cardiac Output During Rest and Strenuous Exercise 50 to 75 mL/100 g muscle/min Most diverts to oxidative portions of muscle at the expense of portions with high glycolytic capacity • For trained individuals, blood redistribution begins just prior to exercise (anticipatory) • Hormonal vascular regulation and local metabolic conditions divert blood flow to active muscles from areas that tolerate compromised blood flow • Blood redistribution among specific tissues occurs primarily during intense exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Blood Flow to the Heart and Brain • The heart and brain cannot tolerate compromised blood supply • At rest, the myocardium uses ~75% of O2 in blood flowing through the coronary circulation • During exercise, the coronary circulation has a four- to fivefold increase in blood flow • Cerebral blood flow increases during exercise by ~25-30% compared with the resting flow Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Cardiac Output and Oxygen Transport: Rest • Arterial blood carries 200 mL O2/L • At rest cardiac output equals 5 L, therefore 1000 mL O2 are available to body • • Resting VO2 averages 250-300 mL/min, allowing 750 mL O2 to return unused to heart • Extra oxygen circulating above resting requirement represents oxygen “in reserve” Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Close Association Between Maximal Cardiac Output and VO2max • Cardiac Output and Oxygen Transport: Physical Activity Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins • • In the untrained, 3200 mL O2 circulates each • Low VO2max corresponds with low maximal cardiac • An increase in maximal cardiac output produces • 5- to 6-L increase in blood flow accompanies each 1-L • minute via a 16-L cardiac output during exercise proportionate increase in capacity to circulate • O2 and increases VO2max output increase in VO2 above rest; this ratio remains unchanged regardless of exercise mode • • High levels of VO2max and maximal cardiac output are distinguishing characteristics for preadolescent and adult endurance athletes • Proportionate increase in maximal cardiac output • accompanies increases in VO2max with endurance training Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 19 Cardiac Output Differences Among Men and Women and Children Close Relationship Between Maximal Cardiac Output and VO2max • • Cardiac output and VO2 linearly relate during graded exercise for boys and girls and men and women • Teenage and •adult women exercise at submaximal VO2 with 5 to 10% larger cardia output than males due to their 10% lower Hb concentration • • Higher submaximal exercise HR in children do not compensate for a smaller SV Produces smaller cardiac output for children - a-vO diff expands to meet the O requirements 2 Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Oxygen Extraction: The a-vO 2 Difference - diff relationship • VO2, cardiac output, and a-vO 2 • expressed in the equation: VO2 = cardiac output × a-vO2diff • Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 2 Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins a-vO 2 Difference During Rest - diff = 5 mL O /dL blood • During rest, a-vO 2 2 perfusing the whole-body tissue–capillary bed 15 mL of O2 (75% of blood’s original O2 content) still remains bound to hemoglobin Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins a-vO 2 Difference During Physical Activity a-vO 2 Difference During Exercise, cont. • Arterial blood O2 content varies little from • Capacity of each deciliter of arterial blood to • Mixed-venous O2 content varies between • Two factors contribute to hemoconcentration 20 mL/dL at rest throughout the exercise intensity range 12 to 15 mL/dL during rest to 2 to 4 mL/dL during maximal exercise - diff results Progressive expansion of a-vO 2 from an increased cellular O2 extraction leading to a reduced venous O2 content Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins carry O2 increases slightly during exercise from increased hemoconcentration from progressive movement of fluid from plasma to interstitial space: Increases in capillary hydrostatic pressure as blood pressure rises Metabolic byproducts of exercise metabolism that osmotically draw fluid from plasma into tissue spaces from plasma Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 20 Changes in a-vO 2 Difference From Rest to Maximal Effort Factors That Affect the a-vO 2 Difference During Physical Activity • Central and peripheral factors interact to increase O2 extraction in active tissue during exercise Large portion of the cardiac output diverts to active tissue Some tissues temporarily decrease blood supply to make more O2 available to tissues in need Exercise training redirects a greater portion of central circulation to active muscle Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Cardiovascular Adjustments to Upper-Body Exercise • • Highest VO2 during arm exercise averages 20 to 30% lower than leg exercise Arm exercise produces lower maximal values for HR and pulmonary ventilation compared to leg exercise Differences relate to smaller muscle mass activated in arm exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Submaximal Oxygen Consumption: Upper-Body Exercise • Higher VO2 during arm exercise at all submaximal power outputs • Two factors that produce additional O2 cost at higher intensities of arm exercise include: 1. Lower mechanical efficiency in upper-body exercise from the additional cost of static muscle actions that do not contribute to external work 2. Recruitment of additional musculature to stabilize torso during arm exercise Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins • VO2 in Arm Exercise Versus Leg Exercise VO2 Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Physiologic Response: Upper-Body Exercise • • Any level of submaximal VO2 or power output with upper-body exercise provides greater physiologic strain than lower-body exercise • Submaximal arm exercise produces higher HR, pulmonary ventilation, perception of effort, BP • Elevated HR response results from two factors: Greater feed-forward stimulation from the brain’s central command to medullary control center Increased feedback stimulation to the medulla from peripheral receptors in active muscle Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 21 Implications of Upper-Body Exercise Summary • Standard submaximal exercise load with the 1. Cardiac output reflects functional capacity of the cardiovascular system: cardiac output = HR x SV upper body produces greater metabolic and physiologic strain than leg exercise • Exercise prescriptions based on running and bicycling do not apply to arm exercise • • Low correlations between VO2max− arm versus • VO 2max− leg exercise do not allow accurate • VO2max predictions for arm exercise based on leg exercise and vice versa • Lack of strong association between two exercise modes verifies the exercise specificity concept Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 2. Cardiac output increases proportionally with effort; from 5 L/min during rest to 20-25 L/min in untrained to 30 to 40 L/min in trained during maximal exercise 3. Differences in maximal cardiac output between athletes and untrained are due to larger SV of athletes 4. SV increases during upright exercise occurs from interaction between greater ventricular filling during diastole and more complete emptying during systole Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Summary, cont. Summary, cont. 5. Sympathetic hormones augment systolic ejection to increase SV during systole 10.Arm exercise generates a 25% lower VO2max than leg exercise 6. Blood flows to specific tissues in proportion to metabolic activity 11.Power output with upper-body exercise provides greater physiologic strain than lower body • exercise at any submaximal VO2 7. Most cardiac output diverts to active muscle during exercise from reduced blood flow to kidneys and splanchnic region • - diff 8. Maximal cardiac output and maximal a-vO 2 • determine VO2max 9. A large cardiac output differentiates endurance athletes from untrained persons Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2015 Wolters Kluwer Health | Lippincott Williams & Wilkins 22