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Conversely, decreasing afterload shifts the curves up and to the left, thereby increasing the stroke volume at any given preload.. Effects of Inotropy on Stroke Volume Effects of Inotrop

Trang 1

of as an adaptive mechanism by which the

ventricle is able to offset the increase in wall

stress that accompanies increased aortic

pres-sure, aortic valve stenosis, or ventricular

dila-tion

Effects of Afterload on Frank-Starling

Curves

An increase in afterload shifts the

Frank-Starling curve down and to the right (Fig

4-15) Therefore, at a given preload (LVEDP

in Figure 4-15), an increase in afterload

de-creases stroke volume Conversely, decreasing

afterload shifts the curves up and to the left,

thereby increasing the stroke volume at any

given preload For reasons discussed later,

changes in afterload result in a subsequent

change in preload so that as the

Frank-Starling curve shifts, the operating point for

the curve shifts diagonally, as shown in Figure

4-15 In the normal heart, changes in

after-load do not have pronounced effects on stroke

volume; however, the failing ventricle is very

sensitive to changes in afterload

Effects of Afterload on the Velocity of Fiber

Shortening (Force-Velocity Relationship)

The decrease in stroke volume that

accompa-nies an increase in afterload is caused by

im-paired emptying of the ventricle The basis

for this is found in the force-velocity rela-tionship of cardiac myocytes The

force-velocity relationship shows how afterload af-fects the velocity of shortening when muscle fibers contract isotonically To illustrate this, a papillary muscle is placed in an in vitro bath and a load is attached to one end (Fig 4-16, left panel) When the muscle contracts, the fiber first generates tension isometrically

(right panel, a to b) until the developed

ten-sion exceeds the load imposed on the muscle When this point is reached, the muscle fiber begins to shorten and the tension remains constant and equal to the load that is being

lifted (b to c) The maximal velocity of

short-ening occurs shortly after the muscle begins

to shorten The muscle continues to shorten until the muscle begins to relax When active

tension falls below the load (point c), the

muscle resumes its resting length (i.e.,

pre-load) (point c) Active tension continues to fall isometrically (c to d) until only the passive tension remains (point d).

If this experiment with the papillary mus-cle were repeated with increasing loads, a de-crease would occur in both the maximal ve-locity of fiber shortening (maximal slope of line) and the degree of shortening, as shown

in Figure 4-17 Plotting the maximal velocity

of shortening against the load that the muscle fiber must shorten against (i.e., the afterload) generates an inverse relationship between ve-locity of shortening and afterload (Fig 4-18)

In other words, the greater the afterload, the slower the velocity of shortening.

To further illustrate the force-velocity rela-tionship, consider the following example If a man holds a 2-pound dumbbell at his side while standing, and then contracts his biceps muscle at maximal effort, the weight will be lifted at a relatively high velocity as the biceps muscle shortens If the weight is increased to

20 pounds, and the weight once again is lifted

at maximal effort, the velocity will be much slower Higher weights further reduce the ve-locity until the weight can no longer be lifted and the contraction of the biceps muscle be-comes isometric The x-intercept in the force velocity diagram (see Fig 4-18) is the point at 10

100

50

20 0

0

LVEDP (mmHg)

B A C

FIGURE 4-15 Effects of changes in afterload on

Frank-Starling curves An increase in afterload shifts the

oper-ating point of the Frank-Starling curve from A to B,

whereas a decrease in afterload shifts the operating

point from A to C Therefore, increased afterload

de-creases stroke volume and inde-creases end-diastolic

pres-sure (preload) The converse also is true.

Trang 2

which the afterload is so great that the muscle

fiber cannot shorten The x-intercept

there-fore represents the maximal isometric force

The y-intercept represents an extrapolated

value for the maximal velocity (Vmax) that

would be achieved if there was no afterload

The value is extrapolated because it cannot be

measured experimentally (a muscle will not

contract in the absence of any load)

It is important to note that a cardiac muscle

fiber does not operate on a single

force-velocity curve (Fig 4-19) If preload is

in-creased, a cardiac muscle fiber will have a

greater velocity of shortening at a given

after-load This occurs because the length-tension relationship requires that as the preload is in-creased, there is an increase in active tension development Once the fiber begins to shorten,

an increased preload with an increase in ten-sion-generating capability causes a greater shortening velocity In other words, increasing the preload enables the muscle to contract faster against a given afterload; this shifts the force-velocity relationship to the right Note that increasing the preload increases the maxi-mal isometric force (x-intercept) as well as the shortening velocity at a given afterload Changes in preload, however, do not alter V

Length Tension

Time

L

T

Preload Resting Maximal

Minimal

∆ L Muscle

Load

a

d

FIGURE 4-16 Cardiac muscle isotonic contractions The left panel shows how muscle length and tension are mea-sured in vitro The lower end of the muscle is attached to a weight (load) that is lifted up from an immovable plat-form as the muscle develops tension and shortens (∆L) A bar attached to the top of the muscle can be moved to

adjust initial muscle length (preload) The right panel shows changes in tension and length during contraction The

periods from a to b and from c to d represent periods of isometric contraction and relaxation, respectively Muscle

shortening (∆L) occurs between b and c, which occurs when the developed tension (∆T) exceeds the load.

Decreasing

Length

Time b

a c = increasing afterload

a

c Preload

FIGURE 4-17 Effects of afterload on myocyte

shorten-ing Increased afterload (curves a to c) decreases the

de-gree of muscle shortening and velocity of shortening at

a given preload.

Afterload (Force)

Maximal Isometric Force Vmax

FIGURE 4-18 Force-velocity relationship Increased af-terload (which requires increased force generation) de-creases velocity of shortening by the muscle fiber The x-intercept represents the maximal isometric force; the y-intercept represents the maximal velocity of

shorten-ing (V max) extrapolated to zero load.

Trang 3

Therefore, an increase in preload on a cardiac

myocyte helps to offset the reduction in velocity

that occurs when afterload is increased.

Effects of Afterload on Pressure-Volume

Loops

Changes in afterload produce secondary

changes in preload, as shown in Figure 4-15

Therefore, afterload and preload are

interde-pendent variables This interdependence can

best be described using pressure-volume loops

(Fig 4-20) If afterload is increased by

increas-ing aortic diastolic pressure, the ventricle has

to generate increased pressure before the

aor-tic valve can open The ejection velocity after

the valve opens will be reduced because

in-creased afterload decreases the velocity of

car-diac fibers shortening, as described by the

force-velocity relationship Because only a

fi-nite period of time exists for electrical and

me-chanical systole, less blood will be ejected

(de-creased stroke volume) so that ventricular

end-systolic volume increases as shown in the

pressure-volume loop The increased

end-sys-tolic volume inside the ventricle will be added

to the venous return, thereby increasing

end-diastolic volume After several beats, a steady

state is achieved in which the increase in

end-systolic volume is greater than the increase in

end-diastolic volume so that the difference

be-tween the two—the stroke volume—is

de-creased (i.e., the width of the pressure-volume

loop is decreased) This increase in preload

secondary to the increase in afterload activates

the Frank-Starling mechanism to partially compensate for the reduction in stroke volume caused by the increase in afterload

Effects of Inotropy on Stroke Volume

Effects of Inotropy on Length-Tension Relationship

Ventricular stroke volume is altered both by changes in preload and afterload, and by

changes in ventricular inotropy (sometimes

referred to as contractility) Changes in ino-tropy are caused by intrinsic cellular mecha-nisms that regulate the interaction between actin and myosin independent of changes in sarcomere length For example, if cardiac

mus-cle is exposed to norepinephrine, it increases active tension development at any initial pload length as shown by the length-tension re-lationship (Fig 4-21) This occurs because the norepinephrine binds to 1-adrenoceptors, in-creasing calcium entry into the cell and cal-cium release by the sarcoplasmic reticulum during contraction (see Chapter 3)

Effects of Inotropy on Force-Velocity Relationship

Changes in inotropy also alter the force-veloc-ity relationship If the inotropic state of the

Afterload (Force)

Increasing Preload a c

FIGURE 4-19 Effects of increasing preload (shift from

curve a to c) on the force-velocity relationship At a

given afterload, increasing the preload increases the

ve-locity of shortening Furthermore, increasing the

pre-load shifts the x-intercept to the right, representing an

increase in isometric force generation.

LV Volume (ml)

Control Loop

Increased Aortic Pressure

FIGURE 4-20 Effects of increased afterload (aortic

pres-sure) on the steady-state left ventricular (LV)

pressure-volume loop Heart rate and inotropy are held constant

in this illustration Increased aortic pressure leads to an

increase in end-systolic volume (ESV), followed by a

sec-ondary, but smaller increase in end-diastolic volume

(EDV) The net effect is a narrower loop and therefore

decreased stroke volume.

Trang 4

myocyte is increased, the force-velocity curve

has a parallel shift up and to the right,

result-ing in an increase in both Vmax and maximal

isometric force (Fig 4-22) The increase in

ve-locity at any given afterload results from the

increased inotropy enhancing force

genera-tion by the actin and myosin filaments and

in-creasing the rate of cross-bridge turnover The

increase in Vmax represents an increased

in-trinsic capability of the muscle fiber to

gener-ate force independent of load

Effects of Inotropy on Pressure-Volume

Loops

The increased velocity of fiber shortening that

occurs with increased inotropy causes an

in-creased rate of ventricular pressure

develop-ment (dP/dt) This increases ejection velocity

and stroke volume and reduces end-systolic

volume, as shown in Figure 4-23 When

ino-tropy is increased, the end-systolic

pressure-volume relationship is shifted to the left and

becomes steeper, because the ventricle can

generate increased pressure at any given

vol-ume The end-systolic pressure-volume

rela-tionship sometimes is used experimentally to

define the inotropic state of the ventricle It is

analogous to the upward shift that occurs in the total tension curve in the length-tension relationship (Fig 4-21) when inotropy in-creases Furthermore, the increased stroke volume leads to a reduction in ventricular end-diastolic volume because less end-systolic volume is available to be added to the incom-ing venous return

Increased Inotropy

Length

Passive Tension

Total Tension

FIGURE 4-21 Effects of increased inotropy on the

length-tension relationship for cardiac muscle

In-creasing inotropy (for example, by stimulating the

car-diac muscle with norepinephrine) shifts the total

tension curve upward, which increases active tension

development (vertical arrows) at any given preload

length.

Afterload (Force)

Increasing Inotropy a c

FIGURE 4-22 Effects of increasing inotropy (parallel

shift from curve a to c) on the force-velocity

relation-ship Increased inotropy increases the velocity of short-ening at any given afterload, and increases V max (y-in-tercept) Furthermore, increased inotropy increases maximal isometric force (x-intercept).

LV Volume (ml)

Increased Inotropy

Control Loop

FIGURE 4-23 Effects of increasing inotropy on the steady state of the left ventricular pressure-volume loop Heart rate and aortic pressure are held constant in this illustration Increased inotropy shifts the end-sys-tolic pressure-volume relationship (see Fig 4-4) up and

to the left, thereby decreasing end-systolic volume

(ESV) A secondary, but smaller decrease in end-diastolic volume (EDV) follows The net effect is an increase in stroke volume (EDV – ESV) LV, left ventricle.

Trang 5

Effects of Inotropy on Frank-Starling

Curves

An increase in inotropy causes the

Frank-Starling curve to shift up and to the left (Fig

4-24) This leads to an increase in stroke

vol-ume along with a reduction in ventricular

pre-load Conversely, a decrease in inotropy (as

occurs in systolic heart failure; see Chapter 9),

shifts the relationship down and to the right,

thereby decreasing stroke volume and

in-creasing preload

Changes in inotropy change the ejection fraction, which is defined as the stroke

vol-ume divided by the end-diastolic volvol-ume A

normal ejection fraction is greater than 0.55

(or 55%) Increasing inotropy increases

ejec-tion fracejec-tion, whereas decreasing inotropy

de-creases ejection fraction Therefore, ejection

fraction often is used as a clinical index for

evaluating the inotropic state of the heart In

heart failure, for example, a decrease in

in-otropy leads to a fall in stroke volume as well

as an increase in preload, thereby decreasing

ejection fraction sometimes to a value less

than 20% Treating a patient in heart failure

with an inotropic drug (e.g.,

agonist or digoxin) shifts the depressed

Frank-Starling curve up and to the left, thereby

in-creasing stroke volume, dein-creasing preload, and increasing ejection fraction

Changes in inotropic state are particularly important during exercise (see Chapter 9) Increases in inotropic state help to maintain stroke volume at high heart rates Increased heart rate alone decreases stroke volume be-cause of reduced time for diastolic filling (de-creased end-diastolic volume) When ino-tropic state increases at the same time, this decreases end-systolic volume to help main-tain stroke volume

Factors Influencing Inotropic State

Several factors influence inotropy (Fig 4-25); the most important of these is the activity of autonomic nerves Sympathetic nerves, by re-leasing norepinephrine that binds to 1 -adrenoceptors on myocytes, are prominent in ventricular and atrial inotropic regulation (see Chapter 3) Parasympathetic nerves (vagal ef-ferents), which release acetylcholine that binds to muscarinic (M2) receptors on my-ocytes (see Chapter 3), have a significant neg-ative inotropic effect in the atria but only a small effect in the ventricles High levels of circulating epinephrine augment sympathetic adrenergic effects via 1-adrenoceptor activa-tion In humans and some other mammalian hearts, an abrupt increase in afterload can

cause a modest increase in inotropy (Anrep effect) by a mechanism that is not fully

un-derstood In addition, an increase in heart rate can cause a small positive inotropic effect

(also termed the Bowditch effect, treppe, or

frequency-dependent activation) This latter phenomenon probably is due to an inability of the Na/K-ATPase to keep up with the sodium influx at higher frequency of action potentials at elevated heart rates, leading to an accumulation of intracellular calcium via the sodium-calcium exchanger (see Chapter 2) Systolic failure that results from cardiomyopa-thy, ischemia, valve disease, arrhythmias, and other conditions is characterized by a loss of intrinsic inotropy In addition to these physio-logic and pathophysio-logic mechanisms, a variety of inotropic drugs are used clinically to increase inotropy in acute and chronic heart failure These drugs include digoxin (inhibits

sar-10

100

50

20 0

0

LVEDP (mmHg)

B

A C

FIGURE 4-24 Effects of changes in inotropy on

Frank-Starling curves Decreased inotropy shifts the operating

point from A to B, which decreases stroke volume and

increases left ventricular end-diastolic pressure (LVEDP).

Increased inotropy causes a shift from point A to C,

which increases stroke volume and decreases LVEDP.

Trang 6

colemmal Na/K-ATPase),

agonists (e.g., dopamine, dobutamine,

epi-nephrine, isoproterenol), and

phosphodi-esterase inhibitors (e.g., milrinone)

Mechanisms of Inotropy

Inotropy can be thought of as a length-inde-pendent activation of the contractile

pro-teins Any cellular mechanism that ultimately

A 67-year-old male is diagnosed with left ventricular failure 4 months following an acute myocardial infarction One of the drugs he is given for treatment acts as a systemic arte-rial vasodilator Using Frank-Starling curves and left ventricular pressure-volume loops, explain how decreasing afterload will improve left ventricular ejection fraction.

A systemic vasodilator reduces afterload on the left ventricle This causes the Starling curve to shift up and to the left from its depressed state (because of the loss of inotropy in failure) (left figure) This shift increases stroke volume and at the same

time reduces preload (end-diastolic pressure) from point A to B in left figure Systemic vasodilation reduces aortic diastolic pressure, which enables the ventricle to eject

sooner, more rapidly, and to a smaller end-systolic volume (right figure) The reduced end-systolic volume leads to a compensatory decrease in end-diastolic volume; how-ever, the reduction in systolic volume will be greater than the reduction in end-diastolic volume so that stroke volume is increased By increasing stroke volume and reducing the end-diastolic volume, the ejection fraction is increased

C A S E 4 - 3

10

100

50

20 0

0

LVEDP (mmHg)

B

A

Non-failing Heart

Failing Heart

LV Volume (ml)

200

100

0

Heart Failure

Arterial Dilator plus

Heart Failure Loop

Effects of an arterial vasodilator on stroke volume

and left ventricular end-diastolic pressure (LVEDP) in

heart failure In heart failure (specifically systolic

fail-ure – see Chapter 9), the Frank-Starling curve shifts

downward because of depressed inotropy Arterial

vasodilation, which reduces afterload on the

ventri-cle, moves the operating point from A to B by

shift-ing the Frank-Starlshift-ing curve upward This leads to an

increase in stroke volume and a decrease in LVEDP

(preload).

Effects of an arterial vasodilator on left ventricular pressure-volume loops Heart failure causes a down-ward shift (reduced slope) of the end-systolic pres-sure-volume relationship This leads to an increase in end-systolic volume, a smaller compensatory in-crease in end-diastolic volume, and reduced stroke volume Reducing arterial pressure decreases afterload on the ventricle, which leads to an increase

in stroke volume This decreases left ventricular systolic volume, and to a smaller extent, end-diastolic volume The net effect is an increase in

stroke volume LV, left ventricle.

Trang 7

alters myosin ATPase activity at a given

sar-comere length alters force generation and

therefore can be considered an inotropic

mechanism Most of the signal transduction

pathways that regulate inotropy involve Ca

(see Chapter 3) Briefly, inotropic state can be

enhanced by (1) increasing Cainflux across

the sarcolemma during the action potential

(via L-type Cachannels); (2) increasing the

release of Ca by the sarcoplasmic

reticu-lum; or (3) sensitizing troponin C to Ca For

example, 1-adrenoceptor activation acting

through Gs-proteins increases cAMP, which

activates protein kinase-A This enzyme can

phosphorylate different intracellular sites to

influence Caentry, Carelease, and Ca

affinity Cardiac glycosides such as digoxin

in-hibit the Na/K-ATPase, leading to an

in-crease in intracellular Caand an increase in

inotropy

MYOCARDIAL OXYGEN

CONSUMPTION

Changes in stroke volume, whether caused by

changes in preload, afterload, or inotropy,

sig-nificantly alter the oxygen consumption of the

heart Changes in heart rate likewise affect

myocardial oxygen consumption The

con-tracting heart consumes a considerable

amount of oxygen because of its need to

re-generate the large amount of ATP hydrolyzed

during contraction and relaxation Therefore,

any change in myocardial function that affects

either the generation of force by myocytes or

their frequency of contraction will alter oxy-gen consumption In addition, even in non-contracting cells, ATP utilized by ion pumps and other transport functions requires oxygen for the resynthesis of ATP

How Myocardial Oxygen Consumption is Determined

Oxygen consumption is defined as the volume

of oxygen consumed per min (e.g., mL

O2/min) and is sometimes expressed per 100 g

of tissue weight (mL O2/min per 100 g) The myocardial oxygen consumption (MVO2) can

be calculated by knowing the coronary blood flow (CBF) and the arterial and venous oxy-gen contents (AO2and VO2) according to the

following equation that uses the Fick Principle:

MVO2 CBF  (AO2 VO2) Myocardial oxygen consumption, there-fore, is equal to the coronary blood flow mul-tiplied by the amount of oxygen extracted from the blood (the arterial-venous oxygen difference) The content of oxygen in blood is usually expressed as mL O2/100 mL blood (or, vol % O2) The oxygen content of arterial blood is normally about 20 mL O2/100 mL blood To calculate the myocardial oxygen consumption in the correct units, mL O2/100

mL blood is converted to mL O2/mL blood; with this conversion, the arterial oxygen con-tent is 0.2 mL O2/mL blood For example, if CBF is 80 mL/min per 100 g, the AO is 0.2

Catecholamines

Heart Rate (Bowditch Effect) Systolic

Failure

Afterload (Anrep Effect)

Sympathetic Activation Parasympathetic

Activation

_

_

Inotropic State (Contractility)

+ +

FIGURE 4-25 Factors regulating inotropy ( ), increased inotropy; (), decreased inotropy.

Eq 4-3

Trang 8

mL O2/mL blood and VO2 is 0.1 mL O2/mL

blood, MVO2 8 mL O2/min per 100 g This

value of myocardial oxygen consumption is

typical for what is found in a heart contracting

at resting heart rates against normal aortic

pressures During heavy exercise, myocardial

oxygen consumption can increase to 70 mL

O2/min per 100 g, or more If contractions are

arrested (e.g., by depolarization of the heart

with a high concentration of potassium

chlo-ride), the myocardial oxygen consumption

de-creases to about 2 mL O2/min per 100g This

value represents the energy costs of cellular

functions not associated with contraction

Therefore, myocardial oxygen consumption

varies considerably depending on the state of

mechanical activity

Although myocardial oxygen consumption

can be calculated as described above,

gener-ally it is not feasible to measure coronary

blood flow and venous oxygen content except

in experimental studies Coronary blood flow

can be measured by placing flow probes on

coronary arteries or a thermodilution catheter

within the coronary sinus Arterial oxygen

content can be taken from a peripheral artery,

but the venous oxygen content has to be

ob-tained from the coronary sinus by inserting a

catheter into the right atrium and then into

the coronary sinus

Indirect indices of myocardial oxygen

con-sumption have been developed to estimate

myocardial oxygen consumption when it is not feasible to measure it Although no index has proven to be satisfactory over a wide range of physiologic conditions, one simple index

sometimes used in clinical studies is the pres-sure-rate product (also called the

double-product) This index can be measured nonin-vasively by multiplying heart rate and systolic arterial pressure (mean arterial pressure sometimes is used instead of systolic arterial pressure) The pressure-rate product assumes that the pressure generated by the ventricle is not significantly different than the aortic pres-sure (i.e., there is no aortic valve stenosis) Experiments have shown that a reasonable correlation exists between changes in the pressure-rate product and myocardial oxygen consumption For example, if arterial pres-sure, heart rate, or both become elevated, oxy-gen consumption will increase

Factors Influencing Myocardial Oxygen Consumption

Part of the difficulty in finding a suitable index

of oxygen consumption is that several factors determine myocyte oxygen consumption, in-cluding frequency of contraction, inotropic state, afterload, and preload (Table 4-1) For example, doubling heart rate approximately doubles oxygen consumption, because myo-cytes are generating twice the number of

ten-In an experimental study, administration of an inotropic drug is found to increase coro-nary blood flow (CBF) from 50 to 150 mL/min and increase the arterial-venous oxygen difference (AO 2 – VO 2 ) from 10 to 14 mL O 2 /100 mL blood Calculate the percent in-crease in myocardial oxygen consumption (MV O 2 ) caused by infusion of this drug.

Myocardial oxygen consumption can be calculated from Equation 4-3, such that

MVO2 CBF  (AO2 VO2) The control oxygen consumption is 50 mL/min times the A-V oxygen difference of 0.1 mL O2/mL blood, which equals 5 mL O2/min Note that the arterial-venous oxygen difference must be converted from mL O2/100 mL blood to mL O2/mL blood The experi-mental oxygen consumption is 150 mL/min times 0.14 mL O2/mL blood, which equals 21

mL O2/min This is a 320% increase in oxygen consumption ([(21 -5)/5] x 100)

P R O B L E M 4 - 3

Trang 9

sion cycles per minute Increasing inotropy

in-creases oxygen consumption because both the

rate of tension development and the

magni-tude of tension are increased, and they both

are associated with increased ATP hydrolysis

and oxygen consumption An increase in

af-terload likewise increases oxygen

consump-tion because it increases the tension that must

be developed by myocytes Increasing stroke

volume by increasing preload (end-diastolic

volume) also increases oxygen consumption

Quantitatively, increased preload has less impact on oxygen consumption than does an

increase in afterload (e.g., aortic pressure) To

understand why, we need to examine the

rela-tionship between wall stress, pressure, and

ra-dius of the ventricle As discussed earlier (see

Equation 4-2), ventricular wall stress () is

proportional to the intraventricular pressure

(P) multiplied by the ventricular internal

ra-dius (r) and divided by the wall thickness (h)

 ∝ Wall stress is related to the tension an indi-vidual myocyte must develop during

contrac-tion to generate a given ventricular pressure

At a given radius and wall thickness, a

my-ocyte must generate increased contractile

force (i.e., wall stress) to develop a higher

pressure The contractile force must be

in-creased even further to generate the same

el-evated pressure if the ventricular radius is

in-creased For example, if the ventricle is

required to generate 50% more pressure than

normal to eject blood because of elevated

aor-P r

h

tic pressure, the wall stress that individual myocytes must generate will be increased by approximately 50% This will increase the oxy-gen consumption of these myocytes by about 50% because changes in oxygen consumption are closely related to changes in wall stress As

a second example, if the radius of the ventri-cle is increased by 50%, the wall stress needed

by the myocytes to eject blood at a normal pressure will be increased by about 50% On the other hand, if the ventricular end-diastolic volume is increased by 50% and the pressure and wall thickness remain unchanged, the wall stress will be increased by only about 14% The reason for this is that a large change

in ventricular volume (V) requires only a small change in radius (r) If we assume that the shape of the ventricle is a sphere, then

V   r3

By rearranging this relationship, we find that

r ∝3V Substituting this into the wall stress equation results in

 ∝ Although no single acceptable model for the shape of the ventricle exists because its shape changes during contraction, a sphere serves as

a convenient model for illustrating why changes in volume have a relatively small af-fect on wall stress and oxygen consumption Using this model, Equation 4-4 shows that in-creasing the end-diastolic volume by 50% (by

a factor of 1.5) represents only a 14% (cube root of 1.5) increase in wall stress at a given ventricular pressure, whereas a 50% increase

in pressure increases wall stress by 50% Therefore, increasing pressure by a given per-centage increases wall stress about four times more than the same change in volume Relating the wall stress equation to oxygen consumption helps to explain why increases in pressure generation have a much greater in-fluence on oxygen consumption than a similar percentage increase in ventricular preload It

is important, however, not to use the wall

P 3V h

4 3

TABLE 4-1 FACTORS INCREASING

MYOCARDIAL OXYGEN

CONSUMPTION

↑ Heart Rate

↑ Inotropy

↑ Afterload

↑ Preload*

*Changes in preload affect oxygen consumption

much less than do changes in the other factors.

Eq 4-4

Trang 10

stress equation to estimate oxygen demands

by the whole heart The reason for this is that

wall stress estimates the tension required by

individual myocytes to generate pressure as

they contract This wall stress, in large part,

determines the oxygen consumption of

indi-vidual myocytes, but oxygen consumption of

the whole heart is the sum of the oxygen

con-sumed by all of the myocytes A

hypertro-phied ventricle with a thicker wall, which has

reduced wall stress, may not have a reduction

in overall oxygen consumption as suggested by

Equation 4-4 In fact, because of its greater

muscle mass, oxygen consumption may be

sig-nificantly increased in a hypertrophied heart,

particularly if its efficiency is impaired by

dis-ease A less efficient heart performs less work

per unit oxygen consumed (i.e., it generates

less pressure and stroke volume)

The concepts described above have

impli-cations for treating patients with coronary

artery disease (CAD) For example, drugs that

decrease afterload, heart rate, and inotropy

are particularly effective in reducing

myocar-dial oxygen consumption and relieving

symp-toms of chest pain (i.e., angina), which results

from inadequate oxygen delivery relative to

the oxygen demands of the myocardium

CAD patients are counseled to avoid activities

such as lifting heavy weights that lead to large

increases in arterial blood pressure In

con-trast, CAD patients are often encouraged to

participate in exercise programs such as

walk-ing that utilize preload changes to augment

cardiac output by the Frank-Starling

mecha-nism It is important to minimize stressful

sit-uations in these patients because stress causes

sympathetic activation of the heart and

vascu-lature that increases heart rate, inotropy, and

afterload, all of which lead to significant

in-creases in oxygen demand by the heart

SUMMARY OF IMPORTANT

CONCEPTS

• The cardiac cycle is divided into two

gen-eral phases: diastole and systole Diastole

refers to the period of time that the

ventri-cles are undergoing relaxation and filling

with blood from the atria Ventricular filling

is primarily passive, although atrial contrac-tion has a variable effect on the final extent

of ventricular filling (end-diastolic volume) Systole, or ventricular contraction, is initi-ated by electrical depolarization of the ven-tricles, which is represented by the QRS complex of the electrocardiogram Ventricular ejection begins when ventricu-lar pressure exceeds the pressure within the outflow tract (aorta or pulmonary artery) and continues until ventricular relaxation causes the ventricular pressures to fall suffi-ciently below the aortic and pulmonary artery pressures to cause the aortic and pul-monic valves to close The volume of blood remaining in the ventricle at the end of ejection is the end-systolic volume

• The first heart sound (S1) originates from closure of the atrioventricular valves (tri-cuspid and mitral) as the ventricles begin to contract The second heart sound (S2) re-sults from the closure of the pulmonic and aortic valves at the end of ventricular sys-tole The third and fourth heart sounds (S3 and S4), when audible, occur during early diastole and atrial contraction, respectively

• Ventricular stroke volume is the difference between the end-diastolic and end-systolic volumes Ventricular ejection fraction is calculated as the stroke volume divided by the end-diastolic volume Ejection fraction

is frequently used in a clinical setting to as-sess the inotropic state of the left ventricle

• Cardiac output is the product of stroke vol-ume and heart rate Normally, cardiac out-put is influenced more by changes in heart rate than by changes in stroke volume; however, impaired regulation of stroke vol-ume can have a significant adverse affect

on cardiac output, as occurs during heart failure

• Ventricular preload is related to the extent

of ventricular filling (end-diastolic volume) and the sarcomere length Preload can be increased by several factors: increased blood volume, augmented venous return, decreased venous compliance (venous con-striction), atrial contraction force, and de-creased heart rate (increases filling time); indirectly, preload can be increased by a

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