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Chemical Energetics of Vascular Smooth Muscle

R. J. Paul

10.1002/cphy.cp020209

Source: Supplement 7: Handbook of Physiology, The Cardiovascular System, Vascular Smooth Muscle

Originally published: 1980

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 null
2 Historical Aspects
3 Energy Metabolism of Vascular Smooth Muscle
3.1 Biochemistry of ATP Synthesis
3.2 Energy Sources
3.3 Respiration of Vascular Tissue
3.4 Aerobic Glycolysis
3.5 ATP Production
3.6 Energy Stores
4 Energy Utilization and Its Relation to Contractility
4.1 Chemical Energetics of Isometric Contraction
4.2 Energetics Under Other Mechanical Conditions
5 Tension Cost of Vascular Smooth Muscle
5.1 Is Tension Cost Dependent on Stimulus?
5.2 Relaxation Speed Theory of Tension Cost
5.3 Rüegg's Approach
5.4 Factors Underlying Low Tension Cost
6 Physiological Correlates
6.1 Metabolic Cost of Vascular Regulation of Circulation
6.2 Physiological Role of Dependence of Force and Metabolism on Tissue Length
6.3 Coordination of Contractility and Energy Metabolism
7 Answers and Questions
Figure 1. Figure 1.

A: relation between active isometric force, ΔPo(l)/ΔPo(lo), and tissue length, l/lo, at various levels of activation. The lo refers to length at which passive force was less than 1% of total force. Heavy curve represents active isometric force–length characteristics of bovine mesenteric vein measured at maximum activation 165. This relation represented for 75% (•), 50% (×), and 25% (▴) of maximum activation defined in terms of percent of maximum isometric force at fixed length. An example of this definition at an arbitrary muscle length is shown by points a–f on vertical bar, representing, 100, 75, 50, 25, 12.5, and 6.25% activation, respectively (arrow points in direction of increasing muscle activation). Muscle lengths corresponding to these levels of activation under fixed‐load conditions (tonus experimental design) are given by the corresponding lettered points on horizontal bar. These points represent idealized characteristics. Discrepancies in length reached by isotonic shortening and length predicted on basis of isometric force‐length relation have been reported for heavy loads 173 and for zero load at low levels of stimulation 203. These discrepancies further complicate the use of relative shortening (tone or tonus) as a definition of muscle activity. Broken line represents passive force–length characteristics. B: ssuprabasal metabolic rate idealized from experiments described in text as a function of 1) isometric force at a given length normalized to the maximum force at that length (ΔPo/ΔPomax shown by heavy line) and corresponding to definition of muscle activity given by vertical bar in A; and 2) length shortened under a given load normalized to the maximum shortening at that load (Δl/Δl max shown by light line) and corresponding to definition of muscle tone as given by horizontal bar in A.
Figure 2. Figure 2.

A: ratio of ATP production attributable to aerobic glycolysis, , to total ATP production, , as a function of molar ratio of oxygen consumption, JO2, to lactate production, Jlac (broken line), and as a function of ratio of carbohydrate utilization attributable to aerobic glycolysis, , to total carbohydrate utilization, (solid line). B: rate of glucose utilization, Jglu, assuming constant ATP utilization, JATP = K, as a function of ratio of ATP production attributable to aerobic glycolysis to total ATP production, . Ordinate is normalized to maximum rate of glucose utilization, which is equal to K/2.5. These relations are calculated on basis of stoichiometry of standard biochemical pathways, assuming that glycogen and glucose are equally likely to be source of carbohydrate. and .
Figure 3. Figure 3.

Record of oxygen tension and isometric force in bovine mesenteric vein for initial stimulation with epinephrine (2.7 μg/ml) after a 2‐h equilibration period, and the second maximal stimulation. Time course of Jo2 (note that time runs right to left) in response to stimulation is very fast, as shown by rapid change in downward slope after the small stimulus artifact due to introduction of stimulant solution. Maximum isometric force is 40 gwt. (Note that sensitivity of initial passive tension is 10 times larger than the following force records.) Top, slope of oxygen tension vs. time trace (mV/min) is given (as determined by straight‐edge fit), and can be converted to Jo2 using 0.05 μmol O2 /mV. Fast vertical rises in oxygen tension trace are due either to partial flushes, which both dilute the stimulant and return oxygen tension to near that of air, or simply to changes in electrical bias to keep the trace on recorder scale.

Adapted from Paul et al. 168
Figure 4. Figure 4.

A: record of isometric force (top numbers) in bovine mesenteric vein during a sequence of decreasing epinephrine concentration after maximal stimulation. Time runs from right to left as shown. Note that initial and final passive‐force measurements were recorded on a scale with fivefold greater sensitivity. Sequence of epinephrine concentrations (μg/ml) right to left is 0, 2.7, 1.6, 0.8, 0.5, 0.3, and <0.01. Periods of oscillatory behavior are seen for some submaximal doses. B: simultaneous record of oxygen electrode output (bottom numbers) during the sequence shown in A. Calibration and vertical rises in O2 tension are as in Figure 3.

From Paul et al. 168
Figure 5. Figure 5.

Relation between oxygen consumption rate and isometric force in bovine mesenteric vein. ○, Measurements taken with increasing epinephrine concentration; •, measurements taken with decreasing epinephrine concentration; bar, basal oxygen consumption rate prior to stimulation. Isometric force is expressed in gram‐weight (1 gwt = 9.8 × 10−3 N).

From Paul et al. 167
Figure 6. Figure 6.

Plot of O2 consumption rate vs. graded isometric tension in bovine mesenteric vein with various stimulants (Epi, epinephrine; Hist, histamine) and the α‐adrenergic blocker dihydroergotamine (Dhe). Regression line shown is for all points; regression lines for categorized subsets of points were not statistically different.

From Paul et al. 168
Figure 7. Figure 7.

A: total lactic acid in bathing solution is shown as a function of time for a series of graded isometric tensions (ΔPo) in bovine mesenteric vein. Arrows indicate additions of epinephrine and corresponding increases in isometric tension. Lines connecting the samples therefore represent lactic acid production rates during 4 intervals of stimulation, which are seen to increase in parallel with isometric tension. B: immediately after a determination of the production rate of basal lactic acid, this vein segment was stimulated to maximum isometric tension (about 50 gwt) with epinephrine (EPI). The two samples after stimulation illustrate that rate of lactic acid production increases abruptly to a new and constant rate. IR (release from isometric) indicates initiation of a lightly loaded contraction, which attains Lmin (minimum contracted length) in about 5 min. Broken line represents extrapolated lactic acid production between the samples taken before and after the active contraction. C: left, plot of data obtained from procedures illustrated in A and B for a single vein segment. Rate of lactic acid production, JLA (Δ), and rate of O2 consumption, Jo2 (○), depend linearly on the graded active isometric tension maintained at resting length. At minimum contracted length (▴, •) where the developed isometric tension with maximal stimulation is small, both JLA and Jo2 are found to be about 20% greater than their respective basal values. Right, JLA and Jo2 are both converted to JATP and summed. A linear dependence on isometric force, ΔPo, and an elevation of JATP at fully shortened length, Lmin, are evident. Note that plot has been expanded; ordinate does not begin at zero.

From Peterson and Paul 172
Figure 8. Figure 8.

Steady‐state measurements of isometric force (Po); rate of lactate production (Jlac), rate of O2 consumption (Jo2), and calculated rate of ATP utilization (JATP) in porcine carotid artery. Measurements were made on a single arterial segment in consecutive order from left to right; each measurement period was approximately 30 min. PSS refers to physiological saline solution, K+‐PSS to a K+ for Na+ substituted PSS, and HIST to histamine stimulation. Lopt refers to optimal length for force development and Lmin to fully shortened length. Please note that scale of left‐hand ordinate is for Jlac and Jo2; right‐hand ordinate refers to JATP and Po.

From Glück and Paul 74
Figure 9. Figure 9.

Typical example of linear relation between suprabasal oxygen consumption rate, ΔJo2, and active isometric force, ΔPo, generated at various tissue lengths is shown for measurements in a single bovine mesenteric vein. Lengths (L) indicated are absolute length in cm; Lo, the unloaded passive length, for this vein was 1.3 cm; fully shortened stimulated length was 0.5 cm. Intercept of ΔJo2−ΔPo regression is statistically different from zero.

From Paul and Peterson 165
Figure 10. Figure 10.

Determinants of vascular smooth muscle energy metabolism under isometric conditions. A: top line represents total metabolic rate as a function of active isometric force, measured under conditions of maximal stimulation at various muscle lengths. Metabolic rates are given in terms of ATP utilization in arbitrary units. The ordinate, however, could be expressed in terms of the rates of oxygen consumption, lactate production, or glucose consumption, as these metabolic parameters show a similar dependence on isometric force. Changes in isometric force are assumed to reflect change in number of available actomyosin interaction sites. Solid horizontal line represents basal metabolism measured in the absence of stimulation. Distance between solid and broken horizontal line represents tension‐independent component of suprabasal metabolism which can be associated with the energy requirements of activation processes. B: top line represents total metabolic rate as a function of active isometric force measured under conditions of fixed muscle length with graded contractions produced at various levels of activation. Difference in slopes of relation between metabolism and force under the different conditions (A, B) can be attributed to different levels of activation at the same force. Broken line represents metabolism associated with activation processes.
Figure 11. Figure 11.

Comparative data (corrected for 20°C) from various smooth and striated muscles. Frog sartorius, •; tortoise rectus femoris, +; bovine heart, ▿; guinea pig taenia coli or uterus, □; oyster tonic adductor, ▵; oyster fast adductor, ▾; Mytilus byssal retractor, ○; pecten striated adductor, ▴; squid mantle, ×. A: relation of maximum shortening velocity, Vo max (in muscle lengths/s), to turnover number of myosin ATPase, e (mol ATP/s per mol myosin), and filament length, l. Abscissa, e/l in arbitrary units. B: contractile tension (kg/cm2) related to number of linkages (cross‐bridges) acting in parallel. Abscissa, product (in arbitrary units) of thick‐filament length, l, and number of cross‐bridges per g muscle, n; actomyosin content (open symbols) or number of actin filaments per cross‐sectional area (closed symbols) were used to deduce n. C: maximum power (product of maximum force and speed) in arbitrary units related to ATPase activity per g muscle. Abscissa, e‐n in μmol Pi/g per s. D: holding economy (s) related to turnover number of myosin ATPase and filament length. Abscissa, l/e in arbitrary units (note that l/e is proportional to l/Vo max from A).
Figure 12. Figure 12.

A: dependence of active isometric force (ΔPo) and suprabasal oxygen consumption rate (ΔJo2) on relative muscle length (L/Lo) is shown for several determinations in a single vein. B: cubic regressions of data from 55 bovine mesenteric veins for normalized active force, ΔPo (solid line), and normalized suprabasal oxygen consumption rate, ΔJo2 (broken line), are shown as a function of muscle length (L) relative to resting length (Lo). Values of normalized suprabasal Jo2 at muscle lengths where active isometric tension upon stimulation is zero are shown; values are not statistically different due to scatter at longer lengths

From Paul and Peterson 165


Figure 1.

A: relation between active isometric force, ΔPo(l)/ΔPo(lo), and tissue length, l/lo, at various levels of activation. The lo refers to length at which passive force was less than 1% of total force. Heavy curve represents active isometric force–length characteristics of bovine mesenteric vein measured at maximum activation 165. This relation represented for 75% (•), 50% (×), and 25% (▴) of maximum activation defined in terms of percent of maximum isometric force at fixed length. An example of this definition at an arbitrary muscle length is shown by points a–f on vertical bar, representing, 100, 75, 50, 25, 12.5, and 6.25% activation, respectively (arrow points in direction of increasing muscle activation). Muscle lengths corresponding to these levels of activation under fixed‐load conditions (tonus experimental design) are given by the corresponding lettered points on horizontal bar. These points represent idealized characteristics. Discrepancies in length reached by isotonic shortening and length predicted on basis of isometric force‐length relation have been reported for heavy loads 173 and for zero load at low levels of stimulation 203. These discrepancies further complicate the use of relative shortening (tone or tonus) as a definition of muscle activity. Broken line represents passive force–length characteristics. B: ssuprabasal metabolic rate idealized from experiments described in text as a function of 1) isometric force at a given length normalized to the maximum force at that length (ΔPo/ΔPomax shown by heavy line) and corresponding to definition of muscle activity given by vertical bar in A; and 2) length shortened under a given load normalized to the maximum shortening at that load (Δl/Δl max shown by light line) and corresponding to definition of muscle tone as given by horizontal bar in A.



Figure 2.

A: ratio of ATP production attributable to aerobic glycolysis, , to total ATP production, , as a function of molar ratio of oxygen consumption, JO2, to lactate production, Jlac (broken line), and as a function of ratio of carbohydrate utilization attributable to aerobic glycolysis, , to total carbohydrate utilization, (solid line). B: rate of glucose utilization, Jglu, assuming constant ATP utilization, JATP = K, as a function of ratio of ATP production attributable to aerobic glycolysis to total ATP production, . Ordinate is normalized to maximum rate of glucose utilization, which is equal to K/2.5. These relations are calculated on basis of stoichiometry of standard biochemical pathways, assuming that glycogen and glucose are equally likely to be source of carbohydrate. and .



Figure 3.

Record of oxygen tension and isometric force in bovine mesenteric vein for initial stimulation with epinephrine (2.7 μg/ml) after a 2‐h equilibration period, and the second maximal stimulation. Time course of Jo2 (note that time runs right to left) in response to stimulation is very fast, as shown by rapid change in downward slope after the small stimulus artifact due to introduction of stimulant solution. Maximum isometric force is 40 gwt. (Note that sensitivity of initial passive tension is 10 times larger than the following force records.) Top, slope of oxygen tension vs. time trace (mV/min) is given (as determined by straight‐edge fit), and can be converted to Jo2 using 0.05 μmol O2 /mV. Fast vertical rises in oxygen tension trace are due either to partial flushes, which both dilute the stimulant and return oxygen tension to near that of air, or simply to changes in electrical bias to keep the trace on recorder scale.

Adapted from Paul et al. 168


Figure 4.

A: record of isometric force (top numbers) in bovine mesenteric vein during a sequence of decreasing epinephrine concentration after maximal stimulation. Time runs from right to left as shown. Note that initial and final passive‐force measurements were recorded on a scale with fivefold greater sensitivity. Sequence of epinephrine concentrations (μg/ml) right to left is 0, 2.7, 1.6, 0.8, 0.5, 0.3, and <0.01. Periods of oscillatory behavior are seen for some submaximal doses. B: simultaneous record of oxygen electrode output (bottom numbers) during the sequence shown in A. Calibration and vertical rises in O2 tension are as in Figure 3.

From Paul et al. 168


Figure 5.

Relation between oxygen consumption rate and isometric force in bovine mesenteric vein. ○, Measurements taken with increasing epinephrine concentration; •, measurements taken with decreasing epinephrine concentration; bar, basal oxygen consumption rate prior to stimulation. Isometric force is expressed in gram‐weight (1 gwt = 9.8 × 10−3 N).

From Paul et al. 167


Figure 6.

Plot of O2 consumption rate vs. graded isometric tension in bovine mesenteric vein with various stimulants (Epi, epinephrine; Hist, histamine) and the α‐adrenergic blocker dihydroergotamine (Dhe). Regression line shown is for all points; regression lines for categorized subsets of points were not statistically different.

From Paul et al. 168


Figure 7.

A: total lactic acid in bathing solution is shown as a function of time for a series of graded isometric tensions (ΔPo) in bovine mesenteric vein. Arrows indicate additions of epinephrine and corresponding increases in isometric tension. Lines connecting the samples therefore represent lactic acid production rates during 4 intervals of stimulation, which are seen to increase in parallel with isometric tension. B: immediately after a determination of the production rate of basal lactic acid, this vein segment was stimulated to maximum isometric tension (about 50 gwt) with epinephrine (EPI). The two samples after stimulation illustrate that rate of lactic acid production increases abruptly to a new and constant rate. IR (release from isometric) indicates initiation of a lightly loaded contraction, which attains Lmin (minimum contracted length) in about 5 min. Broken line represents extrapolated lactic acid production between the samples taken before and after the active contraction. C: left, plot of data obtained from procedures illustrated in A and B for a single vein segment. Rate of lactic acid production, JLA (Δ), and rate of O2 consumption, Jo2 (○), depend linearly on the graded active isometric tension maintained at resting length. At minimum contracted length (▴, •) where the developed isometric tension with maximal stimulation is small, both JLA and Jo2 are found to be about 20% greater than their respective basal values. Right, JLA and Jo2 are both converted to JATP and summed. A linear dependence on isometric force, ΔPo, and an elevation of JATP at fully shortened length, Lmin, are evident. Note that plot has been expanded; ordinate does not begin at zero.

From Peterson and Paul 172


Figure 8.

Steady‐state measurements of isometric force (Po); rate of lactate production (Jlac), rate of O2 consumption (Jo2), and calculated rate of ATP utilization (JATP) in porcine carotid artery. Measurements were made on a single arterial segment in consecutive order from left to right; each measurement period was approximately 30 min. PSS refers to physiological saline solution, K+‐PSS to a K+ for Na+ substituted PSS, and HIST to histamine stimulation. Lopt refers to optimal length for force development and Lmin to fully shortened length. Please note that scale of left‐hand ordinate is for Jlac and Jo2; right‐hand ordinate refers to JATP and Po.

From Glück and Paul 74


Figure 9.

Typical example of linear relation between suprabasal oxygen consumption rate, ΔJo2, and active isometric force, ΔPo, generated at various tissue lengths is shown for measurements in a single bovine mesenteric vein. Lengths (L) indicated are absolute length in cm; Lo, the unloaded passive length, for this vein was 1.3 cm; fully shortened stimulated length was 0.5 cm. Intercept of ΔJo2−ΔPo regression is statistically different from zero.

From Paul and Peterson 165


Figure 10.

Determinants of vascular smooth muscle energy metabolism under isometric conditions. A: top line represents total metabolic rate as a function of active isometric force, measured under conditions of maximal stimulation at various muscle lengths. Metabolic rates are given in terms of ATP utilization in arbitrary units. The ordinate, however, could be expressed in terms of the rates of oxygen consumption, lactate production, or glucose consumption, as these metabolic parameters show a similar dependence on isometric force. Changes in isometric force are assumed to reflect change in number of available actomyosin interaction sites. Solid horizontal line represents basal metabolism measured in the absence of stimulation. Distance between solid and broken horizontal line represents tension‐independent component of suprabasal metabolism which can be associated with the energy requirements of activation processes. B: top line represents total metabolic rate as a function of active isometric force measured under conditions of fixed muscle length with graded contractions produced at various levels of activation. Difference in slopes of relation between metabolism and force under the different conditions (A, B) can be attributed to different levels of activation at the same force. Broken line represents metabolism associated with activation processes.



Figure 11.

Comparative data (corrected for 20°C) from various smooth and striated muscles. Frog sartorius, •; tortoise rectus femoris, +; bovine heart, ▿; guinea pig taenia coli or uterus, □; oyster tonic adductor, ▵; oyster fast adductor, ▾; Mytilus byssal retractor, ○; pecten striated adductor, ▴; squid mantle, ×. A: relation of maximum shortening velocity, Vo max (in muscle lengths/s), to turnover number of myosin ATPase, e (mol ATP/s per mol myosin), and filament length, l. Abscissa, e/l in arbitrary units. B: contractile tension (kg/cm2) related to number of linkages (cross‐bridges) acting in parallel. Abscissa, product (in arbitrary units) of thick‐filament length, l, and number of cross‐bridges per g muscle, n; actomyosin content (open symbols) or number of actin filaments per cross‐sectional area (closed symbols) were used to deduce n. C: maximum power (product of maximum force and speed) in arbitrary units related to ATPase activity per g muscle. Abscissa, e‐n in μmol Pi/g per s. D: holding economy (s) related to turnover number of myosin ATPase and filament length. Abscissa, l/e in arbitrary units (note that l/e is proportional to l/Vo max from A).



Figure 12.

A: dependence of active isometric force (ΔPo) and suprabasal oxygen consumption rate (ΔJo2) on relative muscle length (L/Lo) is shown for several determinations in a single vein. B: cubic regressions of data from 55 bovine mesenteric veins for normalized active force, ΔPo (solid line), and normalized suprabasal oxygen consumption rate, ΔJo2 (broken line), are shown as a function of muscle length (L) relative to resting length (Lo). Values of normalized suprabasal Jo2 at muscle lengths where active isometric tension upon stimulation is zero are shown; values are not statistically different due to scatter at longer lengths

From Paul and Peterson 165
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Article Title: Chemical Energetics of Vascular Smooth Muscle
 

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R. J. Paul. Chemical Energetics of Vascular Smooth Muscle. Compr Physiol 2011, Supplement 7: Handbook of Physiology, The Cardiovascular System, Vascular Smooth Muscle: 201-235. First published in print 1980. doi: 10.1002/cphy.cp020209