(BQ) Part 2 book Millers textbook has contents: Cardiac physiology, gastrointestinal physiology and pathophysiology, hepatic physiology and pathophysiology, renal physiology, pathophysiology, and pharmacology, basic principles of pharmacology,.... and other contents.
Chapter 20 Cardiac Physiology LENA S SUN • JOHANNA SCHWARZENBERGER • RADHIKA DINAVAHI Key Points • The cardiac cycle is the sequence of electrical and mechanical events during the course of a single heartbeat • Cardiac output is determined by the heart rate, myocardial contractility, and preload and afterload • The majority of cardiomyocytes consist of myofibrils, which are rodlike bundles that form the contractile elements within the cardiomyocyte • The basic working unit of contraction is the sarcomere • Gap junctions are responsible for the electrical coupling of small molecules between cells • Action potentials have four phases in the heart • The key player in cardiac excitation-contraction coupling is the ubiquitous second messenger calcium • Calcium-induced sparks are spatially and temporally patterned activations of localized calcium release that are important for excitation-contraction coupling and regulation of automaticity and contractility • β-Adrenoreceptors stimulate chronotropy, inotropy, lusitropy, and dromotropy • Hormones with cardiac action can be synthesized and secreted by cardiomyocytes or produced by other tissues and delivered to the heart • Cardiac reflexes are fast-acting reflex loops between the heart and central nervous system that contribute to the regulation of cardiac function and the maintenance of physiologic homeostasis In 1628, English physician, William Harvey, first advanced the modern concept of circulation with the heart as the generator for the circulation Modern cardiac physiology includes not only physiology of the heart as a pump but also concepts of cellular and molecular biology of the cardiomyocyte and regulation of cardiac function by neural and humoral factors Cardiac physiology is a component of the interrelated and integrated cardiovascular and circulatory physiology This chapter discusses only the physiology of the heart It begins with the physiology of the intact heart The second part of the chapter focuses on cellular cardiac physiology Finally, the various factors that regulate cardiac function are briefly discussed The basic anatomy of the heart consists of two atria and two ventricles that provide two separate circulations in series The pulmonary circulation, a low-resistance and high-capacitance vascular bed, receives output from the right side of the heart, and its chief function is bidirectional gas exchange The left side of the heart provides output for the systemic circulation It functions to deliver oxygen (O2) and nutrients and to remove carbon dioxide (CO2) and metabolites from various tissue beds PHYSIOLOGY OF THE INTACT HEART Understanding of mechanical performance of the intact heart begins with the knowledge of the phases of the cardiac cycle and the determinants of ventricular function CARDIAC CYCLE The cardiac cycle is the sequence of electrical and mechanical events during the course of a single heartbeat Figure 20-1 illustrates (1) the electrical events of a single cardiac cycle represented by the electrocardiogram (ECG) and (2) the mechanical events of a single cardiac cycle represented by left atrial and left ventricular pressure pulses correlated in time with aortic flow and ventricular volume.1 473 474 Reduced ventricular filling—diastalsis Rapid ventricular filling Isovol relax Reduced ejection The cardiac cycle begins with the initiation of the heartbeat Intrinsic to the specialized cardiac pacemaker tissues is automaticity and rhythmicity The sinoatrial (SA) node is usually the pacemaker; it can generate impulses at the greatest frequency and is the natural pacemaker Electrical Events and the Electrocardiogram 32 Electrical events of the pacemaker and the specialized conduction system are represented by the ECG at the body surface (also see Chapters 45 and 47) The ECG is the result of differences in electrical potential generated by the heart at sites of the surface recording The action potential initiated at the SA node is propagated to both atria by specialized conduction tissue that leads to atrial systole (contraction) and the P wave of the ECG At the junction of the interatrial and interventricular septa, specialized atrial conduction tissue converges at the atrioventricular (AV) node, which is distally connected to the His bundle The AV node is an area of relatively slow conduction, and a delay between atrial and ventricular contraction normally occurs at this locus The PR interval represents the delay between atrial and ventricular contraction at the level of the AV node From the distal His bundle, an electrical impulse is propagated through large left and right bundle branches and finally to the Purkinje system fibers, which are the smallest branches of the specialized conduction system Finally, electrical signals are transmitted from the Purkinje system to individual ventricular cardiomyocytes The spread of depolarization to the ventricular myocardium is exhibited as the QRS complex on the ECG Depolarization is followed by ventricular repolarization and the appearance of the T wave on the ECG.2 26 Mechanical Events 120 Aortic valve closes Aortic valve opens 100 Pressure (mm Hg) Rapid ejection Atrial systole Isovol contract PART II: Anesthetic Physiology Aortic pressure 80 Left ventricular pressure Mitral valve closes 60 40 Left atrial pressure 20 Mitral valve opens Aortic blood flow (L/min) Ventricular volume (mL) 38 20 Heart sounds a v Venous pulse c Echocardiogram R P T Q S 0.1 P Ventricular systole 0.2 0.3 0.4 0.5 Time (sec) 0.6 0.7 0.8 Figure 20-1. Electrical and mechanical events during a single cardiac cycle The pressure curves of aortic blood flow, ventricular volume, venous pulse, and electrocardiogram are shown (From Berne RM, Levy MN: The cardiac pump In Cardiovascular physiology, ed St Louis, 2001, Mosby, pp 55-82.) The mechanical events of a cardiac cycle begin with the return of blood to the right and left atria from the systemic and pulmonary circulation, respectively As blood accumulates in the atria, atrial pressure increases until it exceeds the pressure within the ventricle, and the AV valve opens Blood passively flows first into the ventricular chambers, and such flow accounts for approximately 75% of the total ventricular filling.3 The remainder of the blood flow is mediated by active atrial contraction or systole, known as the atrial kick The onset of atrial systole is coincident with depolarization of the sinus node and the P wave While the ventricles fill, the AV valves are displaced upward and ventricular contraction (systole) begins with closure of the tricuspid and mitral valves, which corresponds to the end of the R wave on the ECG The first part of ventricular systole is known as isovolumic or isometric contraction The electrical impulse traverses the AV region and passes through the right and left bundle branches into the Purkinje fibers, which leads to contraction of the ventricular myocardium and a progressive increase in intraventricular pressure When intraventricular pressure exceeds pulmonary artery and aortic pressure, the pulmonic and aortic valves open and ventricular ejection occurs, which is the second part of ventricular systole Ventricular ejection is divided into the rapid ejection phase and the reduced ejection phase During the rapid Chapter 20: Cardiac Physiology ejection phase, forward flow is maximal, and pulmonary artery and aortic pressure is maximally developed In the reduced ejection phase, flow and great artery pressure taper with progression of systole Pressure in both ventricular chambers decreases as blood is ejected from the heart, and ventricular diastole begins with closure of the pulmonic and aortic valves The initial period of ventricular diastole consists of the isovolumic (isometric) relaxation phase This phase is concomitant with repolarization of the ventricular myocardium and corresponds to the end of the T wave on the ECG The final portion of ventricular diastole involves a rapid decrease in intraventricular pressure until it decreases to less than that of the right and left atria, at which point the AV valve reopens, ventricular filling occurs, and the cycle repeats itself VENTRICULAR STRUCTURE AND FUNCTION Ventricular Structure The specific architectural order of the cardiac muscles provides the basis for the heart to function as a pump The ellipsoid shape of the left ventricle (LV) is a result of the laminar layering of spiraling bundles of cardiac muscles (Fig 20-2) The orientation of the muscle bundle is longitudinal in the subepicardial myocardium and circumferential in the middle segment and again becomes longitudinal in the subendocardial myocardium Because of the ellipsoid shape of the LV, regional differences in wall thickness result in corresponding variations in the cross-sectional radius of the left ventricular chamber These regional differences may serve to accommodate the variable loading conditions of the LV.4 In addition, such anatomy allows the LV to eject blood in a corkscrew-type motion beginning from the base and ending at the apex The architecturally complex structure of the LV thus allows maximal shortening of myocytes, which results in increased wall thickness and the generation of force during systole Moreover, release of the twisted LV may provide a suction mechanism for filling of the LV during Cardiac muscle Figure 20-2. Muscle bundles (From Marieb EN: Human anatomy & physiology, ed San Francisco, 2001, Pearson Benjamin Cummings, p 684.) 475 diastole The left ventricular free wall and the septum have similar muscle bundle architecture As a result, the septum moves inward during systole in a normal heart Regional wall thickness is a commonly used index of myocardial performance that can be clinically assessed, such as by perioperative echocardiography or magnetic resonance imaging Unlike the LV, which needs to pump against the higher-pressure systemic circulation, the right ventricle (RV) pumps against a much lower pressure circuit in the pulmonary circulation Consequently, wall thickness is considerably less in the RV In contrast to the ellipsoidal form of the LV, the RV is crescent shaped; as a result, the mechanics of right ventricular contraction are more complex Inflow and outflow contraction is not simultaneous, and much of the contractile force seems to be recruited from interventricular forces of the LV-based septum An intricate matrix of collagen fibers forms a scaffold of support for the heart and adjacent vessels This matrix provides enough strength to resist tensile stretch The collagen fibers are made up of mostly the thick collagen type I fiber, which cross-links with the thin collagen type III fiber, the other major type of collagen.5 Elastic fibers that contain elastin are in close proximity to the collagen fibers They account for the elasticity of the myocardium.6 Ventricular Function Systolic Function The heart provides the driving force for delivering blood throughout the cardiovascular system to supply nutrients and to remove metabolic waste Because of the anatomic complexity of the RV, the traditional description of systolic function is usually limited to the LV Systolic performance of the heart is dependent on loading conditions and contractility Preload and afterload are two interdependent factors extrinsic to the heart that govern cardiac performance Diastolic Function Diastole is ventricular relaxation, and it occurs in four distinct phases: (1) isovolumic relaxation; (2) the rapid filling phase (i.e., the LV chamber filling at variable left ventricular pressure); (3) slow filling, or diastasis; and (4) final filling during atrial systole The isovolumic relaxation phase is energy dependent During the auxotonic relaxation (phases through 4), ventricular filling occurs against pressure It encompasses a period during which the myocardium is unable to generate force, and filling of the ventricular chambers takes place The isovolumic relaxation phase does not contribute to ventricular filling The greatest amount of ventricular filling occurs in the second phase, whereas the third phase adds only approximately 5% of total diastolic volume and the final phase provides 15% of ventricular volume from atrial systole To assess diastolic function, several indices have been developed The most widely used index for examining the isovolumic relaxation phase of diastole is to calculate the peak instantaneous rate of decline in left ventricular pressure (−dP/dt) or the time constant of isovolumic decline in left ventricular pressure (τ) The aortic closing–mitral opening interval and the isovolumic relaxation time and peak rate of left ventricular wall thinning, as determined by echocardiography, have both been used to estimate 476 PART II: Anesthetic Physiology diastolic function during auxotonic relaxation Ventricular compliance can be evaluated by pressure-volume relationships to determine function during the auxotonic phases of diastole.7,8 Many different factors influence diastolic function: magnitude of systolic volume, passive chamber stiffness, elastic recoil of the ventricle, diastolic interaction between the two ventricular chambers, atrial properties, and catecholamines Whereas systolic dysfunction is a reduced ability of the heart to eject, diastolic dysfunction is a decreased ability of the heart to fill Abnormal diastolic function is now being recognized as the predominant cause of the pathophysiologic condition of congestive heart failure.9 Ventricular interactions during systole and diastole are internal mechanisms that function as internal feedback to modulate stroke volume Systolic ventricular interaction involves the effect of the interventricular septum on the function of both ventricles Because the interventricular septum is anatomically linked to both ventricles, it is part of the load against which each ventricle has to work Therefore, any changes in one ventricle will also be present in the other In diastolic ventricular interaction, dilatation of either the LV or RV will have an impact on effective filling of the contralateral ventricle and thereby modify function Preload and Afterload Preload is defined as the ventricular load at the end of diastole, before contraction has started First described by Starling, a linear relationship exists between sarcomere length and myocardial force (Fig 20-3) In clinical practice, surrogate representatives of left ventricular volume such as pulmonary wedge pressure or central venous pressure are used to estimate preload.3 With the development of transesophageal echocardiography, a more direct measure of ventricular volume is available Afterload is defined as systolic load on the LV after contraction has begun Aortic compliance is an additional determinant of afterload.1 Aortic compliance is the ability of the aorta to give way to systolic forces from the ventricle Changes in the aortic wall (dilation or stiffness) can alter aortic compliance and thus afterload Examples of pathologic conditions that alter afterload are aortic stenosis and chronic hypertension Both impede ventricular ejection, thereby increasing afterload Aortic impedance, or aortic pressure divided by aortic flow at that instant, is an accurate means of gauging afterload However, clinical measurement of aortic impedance is invasive Echocardiography can noninvasively estimate aortic impedance by determining aortic blood flow at the time of its maximal increase In clinical practice, the measurement of systolic blood pressure is adequate to approximate afterload, provided that aortic stenosis is not present Preload and afterload can be thought of as the wall stress that is present at the end of diastole and during left ventricular ejection, respectively Wall stress is a useful concept because it includes preload, afterload, and the energy required to generate contraction Wall stress and heart rate are probably the two most relevant indices that account for changes in myocardial O2 demand Laplace’s law states that wall stress (σ) is the product of pressure (P) and radius (R) divided by wall thickness (h)3: σ = P × R/2h The ellipsoid shape of the LV allows the least amount of wall stress such that as the ventricle changes its shape from ellipsoid to spherical, wall stress is increased By using the ratio of the long axis to the short axis as a measure of the ellipsoid shape, a decrease in this ratio would signify a transition from ellipsoid to spherical Thickness of the left ventricular muscle is an important modifier of wall stress For example, in aortic stenosis, afterload is increased The ventricle must generate a much higher pressure to overcome the increased load opposing systolic ejection of blood To generate such high performance, the ventricle increases its wall thickness (left ventricular hypertrophy) By applying Laplace’s law, increased left ventricular wall thickness will decrease wall stress, despite the necessary increase in left ventricular pressure to overcome the aortic stenosis (Fig 20-4).10 In a failing heart, the radius of the LV increases, thus increasing wall stress Frank-Starling Relationship The Frank-Starling relationship is an intrinsic property of myocardium by 200 Stroke volume (mL) Optimal sarcomere length Figure 20-3. Frank-Starling relationship The relationship between sarcomere length and tension developed in cardiac muscles is shown In the heart, an increase in end-diastolic volume is the equivalent of an increase in myocardial stretch; therefore, according to the Frank-Starling law, increased stroke volume is generated Actin Actin Normal resting length 100 Frank-Starling curve Sarcomere length 0 150 Ventricular end-diastolic volume (mL) (EDV) 300 Chapter 20: Cardiac Physiology which stretching of the myocardial sarcomere results in enhanced myocardial performance for subsequent contractions (see Fig 20-3) In 1895, Otto Frank first noted that in skeletal muscle, the change in tension was directly related to its length, and as pressure changed in the heart, a corresponding change in volume occurred.11 In 1914, E H Starling, using an isolated heart-lung preparation as a model, observed that “the mechanical energy set free on passage from the resting to the contracted state is a function of the length of the muscle fiber.”12 If a strip of cardiac muscle is mounted in a muscle chamber under isometric conditions and stimulated at a fixed frequency, then an increase in sarcomere length results in an increase in twitch force Starling concluded that the increased twitch force was the result of a greater interaction of muscle bundles Electron microscopy has demonstrated that sarcomere length (2 to 2.2 μm) is positively related to the amount of actin and myosin cross-bridging and that there is an optimal sarcomere length at which the interaction is maximal This concept is based on the assumption that the increase in cross-bridging is equivalent to an increase in muscle performance Although this theory continues to hold true for skeletal muscle, the force-length relationship in cardiac muscle is more complex When comparing force-strength relationships between skeletal and cardiac muscle, it is noteworthy that the reduction in force is only 10%, even if cardiac muscle is at 80% sarcomere length.11 The cellular basis of the Frank-Starling mechanism is still being investigated and is briefly discussed later in this chapter A common clinical application of Starling’s LV pressure in aortic stenosis 477 law is the relationship of left ventricular end-diastolic volume (LVEDV) and stroke volume The Frank-Starling mechanism may remain intact even in a failing heart.13 However, ventricular remodeling after injury or in heart failure may modify the Frank-Starling relationship Contractility Each Frank-Starling curve specifies a level of contractility, or the inotropic state of the heart, which is defined as the work performed by cardiac muscle at any given end-diastolic fiber Factors that modify contractility will create a family of Frank-Starling curves with different contractility (Fig 20-5).10 Factors that modify contractility are exercise, adrenergic stimulation, changes in pH, temperature, and drugs such as digitalis The ability of the LV to develop, generate, and sustain the necessary pressure for the ejection of blood is the intrinsic inotropic state of the heart In isolated muscle, the maximal velocity of contraction (Vmax) is defined as the maximal velocity of ejection at zero load Vmax is obtained by plotting the velocity of muscle shortening in isolated papillary muscle at varying degrees of force Although this relationship can be replicated in isolated myocytes, Vmax cannot be measured in an intact heart because complete unloading is impossible To measure the intrinsic contractile activity of an intact heart, several strategies have been attempted with varying success Pressure-volume loops, albeit requiring catheterization of the left side of the heart, are currently the best way to determine contractility in an intact heart (Fig 20-6).10 The pressure-volume loop represents an indirect measure of the Frank-Starling relationship between force (pressure) and muscle length (volume) Clinically, the most commonly used noninvasive index of ventricular contractile function is the ejection fraction, which is assessed by echocardiography, angiography, or radionuclide ventriculography Family of Frank-Starling curves Normal—exercise R Wall thickness R Laplace's law Pressure radius Wall stressϭ (Wall thickness) Figure 20-4. In response to aortic stenosis, left ventricular (LV) pressure increases To maintain wall stress at control levels, compensatory LV hypertrophy develops According to Laplace’s law, wall stress = pressure ⋅ radius (R) ÷ (2 × wall thickness) Therefore the increase in wall thickness offsets the increased pressure, and wall stress is maintained at control levels (From Opie LH: Ventricular function In The heart Physiology from cell to circulation, ed Philadelphia, 2004, Lippincott-Raven, pp 355-401.) Ventricular performance Normal LV pressure Running Normal—rest Contractile state of myocardum Walking Heart failure Rest Fatal myocardum depression Ventricular end-diastolic volume (Myocardial stretch) Figure 20-5. A family of Frank-Starling curves is shown A leftward shift of the curve denotes enhancement of the inotropic state, whereas a rightward shift denotes decreased inotropy (From Opie LH: Ventricular function In The heart Physiology from cell to circulation, ed Philadelphia, 2004, Lippincott-Raven, pp 355-401.) 478 PART II: Anesthetic Physiology Ejection fraction = (LVEDV − LVESV) /LVEDV where LVESV is left ventricular end-systolic volume Cardiac Work The work of the heart can be divided into external and internal work External work is expended to eject blood under pressure, whereas internal work is expended within the ventricle to change the shape of the heart and to prepare it for ejection Internal work contributes to inefficiency in the performance of the heart Wall stress is directly proportional to the internal work of the heart.14 External work, or stroke work, is a product of the stroke volume (SV) and pressure (P) developed during ejection of the SV Stroke work = SV × P or (LVEDV − LVESV) × P The external work and internal work of the ventricle both consume O2 The clinical significance of internal work is illustrated in the case of a poorly drained LV during cardiopulmonary bypass Although external work is provided by the roller pump during bypass, myocardial ischemia can still occur because poor drainage of the LV creates tension on the left ventricular wall and increases internal work The efficiency of cardiac contraction is estimated by the following formula8: Cardiac efficiency = External work/Energy equivalent of O2 consumption The corkscrew motion of the heart for the ejection of blood is the most favorable in terms of work efficiency, based on the architecture in a normal LV (with the cardiac muscle bundles arranged so that a circumferentially oriented middle layer is sandwiched by longitudinally Internal work External work End-systolic (ES) PV relationship End-systolic c Ejection Relaxation Aortic valve open b Contraction End-diastolic a e Mitral opening d Filling Ventricular volume Figure 20-6. Pressure-volume (PV) loop Point a depicts the start of isovolumetric contraction The aortic valve opens at point b, and ejection of blood follows (points b→c) The mitral valve opens at point d, and ventricular filling ensues External work is defined by points a, b, c, and d, and internal work is defined by points e, d, and c The PV area is the sum of external and internal work (From Opie LH: Ventricular function In The heart Physiology from cell to circulation, ed Philadelphia, 2004, Lippincott-Raven, pp 355-401.) oriented outer layers) In heart failure, ventricular dilation reduces cardiac efficiency because it increases wall stress, which in turn increases O2 consumption.11 Heart Rate and Force-Frequency Relationship In isolated cardiac muscle, an increase in the frequency of stimulation induces an increase in the force of contraction This relationship is termed the treppe, which means staircase in German, and is the phenomenon or the force-frequency relationship.8,15 At between 150 and 180 stimuli per minute, maximal contractile force is reached in an isolated heart muscle at a fixed muscle length Thus an increased frequency incrementally increases inotropy, whereas stimulation at a lower frequency decreases contractile force However, when the stimulation becomes extremely rapid, the force of contraction decreases In the clinical context, pacing-induced positive inotropic effects may be effective only up to a certain heart rate, based on the force-frequency relationship In a failing heart, the force-frequency relationship may be less effective in producing a positive inotropic effect.8 CARDIAC OUTPUT Cardiac output is the amount of blood pumped by the ˙ ) and is determined by four facheart per unit of time (Q tors: two factors that are intrinsic to the heart—heart rate and myocardial contractility—and two factors that are extrinsic to the heart but functionally couple the heart and the vasculature—preload and afterload Heart rate is defined as the number of beats per minute and is mainly influenced by the autonomic nervous system Increases in heart rate escalate cardiac output as long as ventricular filling is adequate during diastole Contractility can be defined as the intrinsic level of contractile performance that is independent of loading conditions Contractility is difficult to define in an intact heart because it cannot be separated from loading conditions.8,15 For example, the Frank-Starling relationship is defined as the change in intrinsic contractile performance, based on changes in preload Cardiac output in a living organism can be measured with the Fick principle (a schematic depiction is illustrated in Fig 20-7).1 The Fick principle is based on the concept of conservation of mass such that the O2 delivered from pulmonary venous blood (q3) is equal to the total O2 delivered to pulmonary capillaries through the pulmonary artery (q1) and the alveoli (q2) The amount of O2 delivered to the pulmonary capillaries by way of the pulmonary arteries (q1) equals total ˙ ) times the O concenpulmonary arterial blood flow (Q tration in pulmonary arterial blood (CpaO2): ˙ × CpaO q1 = Q The amount of O2 carried away from pulmonary venous blood (q3) is equal to total pulmonary venous ˙ ) times the O concentration in pulmonary blood flow (Q venous blood (Cpvo2): ˙ × CpvO q3 = Q The pulmonary arterial O2 concentration is the mixed systemic venous O2, and the pulmonary venous O2 concentration is the peripheral arterial O2 O2 consumption Chapter 20: Cardiac Physiology is the amount of O2 delivered to the pulmonary capillaries from the alveoli (q2) Because q1 + q2 = q3, ( ( ) ) ˙ CpaO + q = Q ˙ CpvO Q 2 ( ( ) ) ˙ CpvO − Q ˙ CpaO q2 = Q 2 ( ) ˙ Cpvo − CpaO q2 = Q 2 ( ) ˙ = q / CpvO − CpaO Q 2 Thus if the CpaO2, CpvO2, and O2 consumption (q2) are known, then the cardiac output can be determined The indicator dilution technique is another method for determining cardiac output also based on the law of conservation of mass The two most commonly used indicator dilution techniques are the dye dilution and the thermodilution methods Figure 20-8 illustrates the principles of the dye dilution method.1 479 and (3) extracellular connective tissue A group of cardiomyocytes with its connective tissue support network or extracellular matrix make up a myofiber (Fig 20-9) Adjacent myofibers are connected by strands of collagen The extracellular matrix is the synthetic product of fibroblasts and is made up of collagen, which is the main determinant of myocardial stiffness, and other major matrix proteins One of the matrix proteins, elastin, is the chief constituent of elastic fibers The elastic fibers account for, in part, the elastic properties of the myocardium.6 Other matrix proteins include the glycoproteins or proteoglycans and matrix metalloproteinases Proteoglycans are proteins with short sugar chains, and they include heparan sulfate, chondroitin, fibronectin, and laminin Matrix metalloproteins are enzymes that degrade collagen and other extracellular proteins The balance between the accumulation of extracellular matrix proteins by synthesis and Mixer Q CELLULAR CARDIAC PHYSIOLOGY A CELLULAR ANATOMY From pulmonary artery To pulmonary veins Terminal bronchiole Photocell Densitometer t1 Alveoli t2 Time Figure 20-8. Illustration demonstrates the principle of determining cardiac output with the indicator dilution technique This model assumes that there is no recirculation A known amount of dye (q) ˙ (mL/min) A mixed is injected at point A into a stream flowing at Q sample of the fluid flowing past point B is withdrawn at a constant rate through a densitometer The change in dye concentration over time is depicted in a curve Flow may be measured by dividing the amount of indicator injected upstream by the area under the downstream concentration curve (From Berne RM, Levy MN: The cardiac pump In Cardiovascular physiology, ed St Louis, 2001, Mosby, pp 55-82.) O2 Consumption 250 mL O2 /min q1 q mg dye injected Dye concentration at point B At the cellular level, the heart consists of three major components: (1) cardiac muscle tissue (contracting cardiomyocytes), (2) conduction tissue (conducting cells), B Lamp q2 [O2 ] pa 0.15 mL O2 /mL blood q3 [O2 ] pv 0.20 mL O2 /mL blood q1ϩq2ϭq3 Figure 20-7. Illustration demonstrates the principle of determination of cardiac output according to the Fick formula If the oxygen (O2) concentration in pulmonary arterial blood (CpaO2), the O2 concentration of the pulmonary vein (CpvO2), and the O2 consumption are known, then cardiac output can be calculated pa, Pulmonary artery; pv, pulmonary vein (From Berne RM, Levy MN: The cardiac pump In Cardiovascular physiology, ed St Louis, 2001, Mosby, pp 55-82.) Myofibrils Figure 20-9. Organization of cardiomyocytes Fifty percent of cardiomyocyte volume is made up of myofibrils; the remainder consists of mitochondria, nucleus, sarcoplasmic reticulum, and cytosol 480 PART II: Anesthetic Physiology Intercalated disks Mitochondrion Cardiac muscle cell Gap junction Nucleus Sarcolemma Desmosome Figure 20-10. The sarcolemma that envelops cardiomyocytes becomes highly specialized to form the intercalated disks where ends of neighboring cells are in contact The intercalated disks consist of gap junctions and spot and sheet desmosomes their breakdown by matrix metalloproteins contributes to the mechanical properties and function of the heart.6 CARDIOMYOCYTE STRUCTURE AND FUNCTION Individual contracting cardiomyocytes are large cells between 20 μm (atrial cardiomyocytes) and 140 μm (ventricular cardiomyocytes) in length Approximately 50% of the cell volume in a contracting cardiomyocyte is made up of myofibrils, and the remainder consists of mitochondria, nucleus, sarcoplasmic reticulum (SR), and cytosol The myofibril is the rodlike bundle that forms the contractile elements within cardiomyocytes Within each contractile element are contractile proteins, regulatory proteins, and structural proteins Contractile proteins make up approximately 80% of the myofibrillar protein, with the remainder being regulatory and structural proteins.16,17 The basic unit of contraction is the sarcomere (see discussion under “Contractile Elements” later in this chapter) The sarcolemma, or the outer plasma membrane, separates the intracellular and extracellular space It surrounds the cardiomyocyte and invaginates into the myofibrils through an extensive tubular network known as transverse tubules or T tubules, and it also forms specialized intercellular junctions between cells.18,19 Transverse or T tubules are in close proximity to an intramembranous system and the SR, which plays an important role in the calcium (Ca2+) metabolism that is critical in the excitation-contraction coupling (ECC) of the cardiomyocyte The SR can be further divided into the longitudinal (or network) SR and the junctional SR The longitudinal SR is involved in the uptake of Ca2+ for the initiation of relaxation The junctional SR contains large Ca2+-release channels (ryanodine receptors [RyRs]) that release SR Ca2+ stores in response to depolarizationstimulated Ca2+ influx through the sarcolemmal Ca2+ channels The RyRs are not only Ca2+-release channels, but they also form the scaffolding proteins that anchor many of the key regulatory proteins.20 Mitochondria are immediately found beneath the sarcolemma, wedged between myofibrils within the cell They contain enzymes that promote the generation of adenosine triphosphate (ATP), and they are the energy powerhouse for the cardiomyocyte In addition, mitochondria can also accumulate Ca2+ and thereby contribute to the regulation of the cytosolic Ca2+ concentration Nearly all of the genetic information is found within the centrally located nucleus The cytosol is the fluid-filled microenvironment within the sarcolemma, exclusive of the organelles and the contractile apparatus and proteins Cardiac muscle cells contain three different types of intercellular junctions: gap junctions, spot desmosomes, and sheet desmosomes (or fasciae adherens) (Fig 20-10).18,21 Gap junctions are responsible for electrical coupling and the transfer of small molecules between cells, whereas desmosome-like junctions provide mechanical linkage The adhesion sites formed by spot desmosomes anchor the intermediate filament cytoskeleton of the cell; those formed by the fasciae adherens anchor the contractile apparatus Gap junctions consist of clusters of plasma membrane channels directly linking the cytoplasmic compartments of neighboring cells Gap junction channels are constructed from connexins, a multigene family of conserved proteins The principal connexin isoform of the mammalian heart is connexin 43; other connexins, notably connexins 40, 45, and 37, are also expressed but in smaller quantities.20,21 The conducting cardiomyocytes, or Purkinje cells, are cells specialized for conducting propagated action potentials These cells have a low content of myofibrils and a prominent nucleus, and they contain an abundance of gap junctions Cardiomyocytes can be functionally separated into (1) the excitation system, (2) the ECC system, and (3) the contractile system Excitation System The cellular action potential originating in the specialized conduction tissue is propagated to individual cells where it initiates the intracellular event that leads to the contraction of the cell through the sarcolemmal excitation system Chapter 20: Cardiac Physiology Transmembrane potential, mV +25 –25 –50 –75 –100 Na+ influx Ca+ influx K+ efflux Na+ efflux K+ influx Figure 20-11. Phases of cellular action potentials and major associated currents in ventricular myocytes The initial phase (0) spike and overshoot (1) are caused by a rapid inward sodium (Na+) current, the plateau phase (2) by a slow calcium (Ca2+) current through L-type Ca channels, and repolarization (phase 3) by outward potassium (K+) currents Phase 4, the resting potential (Na+ efflux, K+ influx), is maintained by Na+-K+-adenosine triphosphatase (ATPase) The Na+-Ca2+ exchanger is mainly responsible for extrusion of Ca2+ In specialized conduction system tissue, spontaneous depolarization takes place during phase until the voltage resulting in opening of the Na channel is reached (From LeWinter MM, Osol G: Normal physiology of the cardiovascular system In Fuster V, Alexander RW, O’Rourke RA, editors: Hurst’s the heart, ed 10 New York, 2001, McGraw-Hill, pp 63-94.) 481 L-type Ca2+ channels and the efflux of K+ through several K+ channels—the inwardly rectifying ik, the delayed rectifier ik1, and ito Repolarization (phase 3) is brought about when an efflux of K+ from the three outward K+ currents exceeds the influx of Ca2+, thus returning the membrane to the resting potential Very little ionic flux occurs during diastole (phase 4) in a fast-response action potential In contrast, during diastole (phase 4), pacemaker cells that show slow-response action potentials have the capability of spontaneous diastolic depolarization and generate the automatic cardiac rhythm Pacemaker currents during phase are the result of an increase in the three inward currents and a decrease in the two outward currents The three inward currents that contribute to spontaneous pacemaker activity include two carried by Ca2+, iCaL and iCaT, and one that is a mixed cation current, If.22 The two outward currents are the delayed rectifier K+ current, ik, and the inward rectifying K+ current, ik1 When compared with the fast-response action potential, phase is much less steep, phase is absent, and phase is indistinct from phase in the slow-response action potential.23 In SA node cells, the pacemaker If current is the principal determinant of duration diastolic depolarization, and it is encoded by four members of the hyperpolarizationactivated cyclic nucleotide-gated gene (HCN1-4) family.24 During the cardiac action potential, movement of Ca2+ into the cell and Na+ out of the cell creates an ionic imbalance The Na+-Ca2+ exchanger restores cellular ionic balance by actively transporting Ca2+ out of the cell against a concentration gradient while moving Na+ into the cell in an energy-dependent manner Excitation-Contraction Coupling Action Potential Ion fluxes across plasma membranes result in depolarization (attaining a less negative membrane potential) and repolarization (attaining a more negative membrane potential) They are mediated by membrane proteins with ion-selective pores Because these ion channel proteins open and close the pores in response to changes in membrane potential, the channels are voltage gated In the heart, sodium (Na+), potassium (K+), Ca2+, and chloride (Cl−) channels contribute to the action potential The types of action potential in the heart can be separated into two categories: (1) fast-response action potentials, which are found in the His-Purkinje system and atrial or ventricular cardiomyocytes; and (2) slowresponse action potentials, which are found in the pacemaker cells in the SA and AV nodes A typical tracing of an action potential in the His-Purkinje system is depicted in Figure 20-11.8 The electrochemical gradient for K+ across the plasma membrane is the determinant for the resting membrane potential Mostly as a result of the influx of Na+, the membrane potential becomes depolarized, which leads to an extremely rapid upstroke (phase 0) As the membrane potential reaches a critical level (or threshold) during depolarization, the action potential is propagated The rapid upstroke is followed by a transient repolarization (phase 1) Phase is a period of brief and limited repolarization that is largely attributable to the activation of a transient outward K+ current, ito The plateau phase (phase 2) occurs with a net influx of Ca2+ through Structures that participate in cardiac ECC include the sarcolemma, transverse tubules, SR, and myofilaments (Fig 20-12, A).25 The process of ECC begins with depolarization of the plasma membrane and spread of electrical excitation along the sarcolemma of cardiomyocytes The ubiquitous second messenger Ca2+ is the key player in cardiac ECC (see Fig 20-12, B).23 Cycling of Ca2+ within the structures that participate in ECC initiates and terminates contraction Activation of the contractile system depends on an increase in free cytosolic Ca2+ and its subsequent binding to contractile proteins Ca2+ enters through plasma membrane channels concentrated at the T tubules, and such entry through L-type Ca2+ channels (dihydropyridine receptors) triggers the release of Ca2+ from the SR.26 This evokes a Ca2+ spark Ca2+ sparks are considered to be the elementary Ca2+ signaling event of ECC in heart muscle A Ca2+ spark occurs with the opening of a cluster of SR RyRs to release Ca2+ in a locally regenerative manner It, in turn, activates the Ca2+-release channels and induces further release of Ca2+ from subsarcolemmal cisternae in the SR and thus leads to a large increase in intracellular Ca2+ (iCa2+) These spatially and temporally patterned activations of localized Ca2+ release, in turn, stimulate myofibrillar contraction The increase in iCa2+, however, is transient inasmuch as Ca2+ is removed by (1) active uptake by the SR Ca2+ pump adenosine triphosphatase (ATPase), (2) extrusion of Ca2+ from the cytosol by the Na+-Ca2+ exchanger, and (3) binding of Ca2+ to proteins.27 Ca2+ sparks have also been implicated 482 PART II: Anesthetic Physiology Extracellular space Plasma Ca2+-ATPase Na+-Ca+ exchanger membrane Na Sodium pump B1 Na B2 T tubule Cytosol L-type calcium channel Calcium release channel Sarcotubular network Subsarcolemmal cisterna Calsequestrin Extracellular space Sarcolplasmic reticulum Phospholamban G C D A1 Thick filament Actin A A SERCA 2A Thin filament E Mitochondria Myosin Troponin Z line cross-bridge Contractile proteins F H B Figure 20-12. A, Diagram depicts the components of cardiac excitation-contraction coupling Calcium pools are noted in bold letters B, Extracellular (arrows A, B1, B2) and intracellular calcium flux (arrows C, D, E, F, and G) are shown The thickness of the arrows indicates the magnitude of the calcium flux, and the vertical orientations describe their energetics: downward-pointing arrows represent passive calcium flux, whereas upward-pointing arrows represent energy-dependent calcium transport Calcium entering the cell from extracellular fluid through L-type calcium channels triggers the release of calcium from the sarcoplasmic reticulum Only a small portion directly activates the contractile proteins (arrow A1) Arrow B1 depicts active transport of calcium into extracellular fluid by means of the plasma membrane calcium adenosine triphosphatase (Ca2+-ATPase) pump and the sodium-calcium (Na+-Ca2+) exchanger Sodium that enters the cell in exchange for calcium (dashed line) is pumped out of the cytosol by the sodium pump SR regulates calcium efflux from the subsarcolemmal cisternae (arrow C) and calcium uptake into the sarcotubular network (arrow D) Arrow G represents calcium that diffuses within the SR Calcium binding to (arrow E) and dissociation from (arrow F) high-affinity calcium-binding sites of troponin C activate and inhibit interactions of the contractile proteins Arrow H depicts movement of calcium into and out of mitochondria to buffer the cytosolic calcium concentration SERCA 2A, Sarcoplasmic/endoplasmic reticulum Ca2+-ATPase (From Katz AM: Calcium fluxes In Physiology of the heart, ed Philadelphia, 2001, Lippincott-Raven, pp 232-233.) in pathophysiologic diseases such as hypertension, cardiac arrhythmias, heart failure, and muscular dystrophy.28-30 The SR provides the anatomic framework and is the major organelle for the cycling of Ca2+ It is the depot for iCa2+ stores The cyclic release plus reuptake of Ca2+ by the SR regulates the cytosolic Ca2+ concentration and couples excitation to contraction The physical proximity between L-type Ca2+ channels and RyRs at the SR membrane makes Ca2+-induced Ca2+ release to occur easily The foot region of the RyR is the part that extends from the SR membrane to the T tubules, where the L-type Ca2+ channels are located.17,27,31 The SR is also concerned with the reuptake of Ca2+ that initiates relaxation or terminates contraction The sarcoplasmic/endoplasmic reticulum Ca2+-ATPase (SERCA) pump is the ATP-dependent pump that actively pumps the majority of the Ca2+ back into the SR after its release SERCA makes up close to 90% of all of the SR proteins and is inhibited by the phosphoprotein, phospholamban, at rest Phospholamban is an SR membrane protein that is active in the dephosphorylated form Phosphorylation by a variety of kinases as a result of β-adrenergic stimulation or other stimuli inactivates phospholamban and releases its inhibitory action on SERCA Positive feedback ensues and leads to further phospholamban phosphorylation and greater SERCA activity Active reuptake of Ca2+ by SERCA then promotes relaxation.17,27,31 Once taken up into the SR, Ca2+ is stored until it is released during the next cycle Calsequestrin and calreticulin are two storage proteins in the SR Calsequestrin is a highly charged protein located in the cisternal component of the SR near the T tubules Because it lies close to the Ca2+-release channels, the stored Ca2+ can be quickly discharged for release once the Ca2+-release channels are stimulated Cytosolic Ca2+ can also be removed by extrusion through the sarcolemmal Ca2+ pump and the activity of the Na+-Ca2+ exchanger The protein, calmodulin, is an important sensor and regulator of iCa2+.19 Chapter 39: Anesthetic Implications of Concurrent Diseases 1221 recurrent supraventricular and ventricular tachycardia, causes thyroid dysfunction as a result of the large amount of iodine in its structure (see the section on thyroid disorders earlier in this chapter), as well as peripheral neuropathy, and has been associated with hypertension, bradyarrhythmias, and reduced cardiac output during anesthesia.465 The drug has a half-life of 29 days, and its pharmacologic effects persist for more than 45 days after discontinuance.466 possibility of fluorine-associated renal damage after enflurane administration.467 Appropriate antibiotic prophylaxis for surgery requires a knowledge of the probability of infection for that type of surgical procedure and, if the incidence of infection warrants, the use of a drug regimen directed against the most likely infecting organisms.468 ANTIBIOTICS Medications for glaucoma include two organophosphates: echothiophate and isoflurophate (see also Chapter 84) These drugs inhibit serum cholinesterase, which is responsible for the hydrolysis and inactivation of succinylcholine and ester-type local anesthetics such as procaine, chloroprocaine, and tetracaine (see also Chapters 34 and 36).469,470 These ester-type local anesthetics should be avoided in patients treated with eye drops containing organophosphate Table 39-16 lists other medications related to anesthesia and their side effects (from the National Registry for Drug-Induced Ocular Side Effects, Many antibacterial drugs are nephrotoxic or neurotoxic, or both, and many prolong neuromuscular blockade (see also Chapters 34 and 35).458-464 The only antibiotics devoid of neuromuscular effects appear to be penicillin G and the cephalosporins.463 Most enzyme-inducing drugs not increase the metabolism of enflurane or isoflurane However, isoniazid induces the microsomal enzymes responsible for the metabolism of at least enflurane and thereby increases the MEDICATIONS FOR GLAUCOMA TABLE 39-16 COMMON OPHTHALMOLOGIC DRUGS AND THEIR ANESTHETICALLY IMPORTANT INTERACTIONS Drug (Trade Name) Toxicities and Specific Treatments Glaucoma: Primary Goal Is to Reduce IOP By Miotics and epinephrine: increase outflow of aqueous humor β-Blockade and carbonic anhydrase inhibitors: reduce production of aqueous humor Osmotic drugs: transiently decrease volume Miotics Parasympathomimetics Pilocarpine (Adsorbocarpine, Isopto Carpine, Pilocar, Pilocel) Carbachol Acetylcholinesterase Inhibitors Physostigmine Demecarium Isoflurophate (Floropryl) Echothiophate (Echodide, Phospholine) Tox: Hypersalivation, sweating, N/V, bradycardia, hypotension, bronchospasm, CNS effects, coma, respiratory arrest, death Rx: Atropine, pralidoxime (Protopam) Ix: Succinylcholine—prolonged apnea (drugs must be discontinued wk before) Epinephrine (Epitrate, Murocoll, Mytrate, Epifrin, Glaucon, Epinal, Eppy) Tox: (rare) Tachycardia, PVCs, HTN, headache, tremors Ix: Avoid drugs that sensitize to catecholamines (e.g., halothane) β-Blockers Timolol (Timoptic) Betaxolol (Betoptic) Levobunolol (Betagan) Tox: β-Blockade with bradycardia, exacerbation of asthma, CNS depression, lethargy, confusion Synergy noted with systemic drugs Carbonic Anhydrase Inhibitors Acetazolamide (Diamox) Dichlorphenamide (Daranide, Oratrol) Ethoxzolamide (Cardrase, Ethamide) Methazolamide (Neptazane) Tox: Anorexia, GI disturbances, “general miserable feeling” and malaise, paresthesias, diuresis, hypokalemia (transient), renal colic and calculi, hyperuricemia, thrombocytopenia, aplastic anemia, acute respiratory failure in patients with COPD Osmotic Drugs Glycerin (Glyrol, Osmoglyn) Isosorbide (Ismotic) Urea (Urevert, Ureaphil) Mannitol (Osmitrol) Intraocular acetylcholine (Miochol) Tox: Dehydration, hyperglycemia, nonketotic hyperosmolar coma (rare); fatalities with mannitol secondary to CHF or intracranial bleeding; urea may cause thrombosis Tox: Hypotension, bradycardia Rx: Atropine Mydriatics and Cycloplegics: Provide Pupillary Dilatation and Paralysis of Accommodation Anticholinergics block muscarinic receptors; paralyzing in iris α-Adrenergics contract the dilator of the iris Continued 1222 PART IV: Anesthesia Management TABLE 39-16 COMMON OPHTHALMOLOGIC DRUGS AND THEIR ANESTHETICALLY IMPORTANT INTERACTIONS—cont’d Drug (Trade Name) Toxicities and Specific Treatments Anticholinergics Atropine (Atropisol, Bufopto, Isopto Atropine) Cyclopentolate, alone (Cyclogyl) or with phenylephrinehomatropine (Cyclomydril) Homatropine (Homatrocel, Isopto Homatropine) Scopolamine (Isopto Hyoscine, Murocoll 19) Tropicamide (Midriacyl) Tox: Dry mouth, flushing, thirst, tachycardia, seizure, hyperactivity, transient psychosis, rare coma, and death Rx: Physostigmine β-Adrenergics Phenylephrine (Efricel, Mydfrin, Neo-Synephrine) Hydroxyamphetamine (Paredrine) Tox: Tachycardia, HTN, PVCs, myocardial ischemia, agitation Modified from the National Registry for Drug-Induced Ocular Side Effects, Portland, Ore., Oregon Health Sciences University CHF, Congestive heart failure; CNS, central nervous system; COPD, chronic obstructive pulmonary disease; GI, gastrointestinal; HTN, hypertension; IOP, intraocular pressure; Ix, interaction; N/V, nausea and vomiting; PVCs, premature ventricular contractions; Rx, treatment; Tox, toxicity Oregon Health Sciences University, 3181 SW Sam Jackson Park Road, Portland, Ore 97201; 503-279-8456) Complete references available online at expertconsult.com Acknowledgment The editors and publisher would like to thank Drs Michael F Roizen and Lee A Fleisher for contributing a chapter on this topic to the seventh edition of this work It has served as the foundation for the current chapter References Wei JY: N Engl J Med 327:1735, 1992 Fleisher LA, Eagle KA: N Engl J Med 345:1677, 2001 Goldman L, et al: N Engl J Med 297:845, 1977 Fleisher LA, et al: J Am Coll Cardiol 50:159, 2007 Diabetes Control and Complications Trial (DCCT)/Epidemiology of Diabetes Interventions and Complications Research Group: N Engl J Med 342:381, 2000 U.K Prospective Diabetes Study Group: BMJ 317:703, 1998 Albacker T, et al: Ann Thorac Surg 86:20, 2008 Krinsley JS: Mayo Clin Proc 79:992, 2004 Advance Collaborative Group: N Engl J Med 358:2560, 2008 10 Van 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