Am J Physiol Regulatory Integrative Comp Physiol 279: R1849–R1855, 2000.

Hemodynamic responses to static and dynamic muscle contractions at equivalent workloads
JASON W. DANIELS,1 CHARLES L. STEBBINS,1 AND JOHN C. LONGHURST2 1 Division of Cardiovascular Medicine, Department of Internal Medicine, University of California, Davis, Davis 95616–8634; and 2Department of Medicine, University of California, Irvine, Irvine, California 92888
Received 24 March 2000; accepted in final form 19 July 2000

Daniels, Jason W., Charles L. Stebbins, and John C. Longhurst. Hemodynamic responses to static and dynamic muscle contractions at equivalent workloads. Am J Physiol Regulatory Integrative Comp Physiol 279: R1849–R1855, 2000.—We tested the hypothesis that static contraction causes greater reflex cardiovascular responses than dynamic contraction at equivalent workloads [i.e., same tension-time index (TTI), holding either contraction time or peak tension constant] in chloralose-anesthetized cats. When time was held constant and tension was allowed to vary, dynamic contraction of the hindlimb muscles evoked greater increases (means Ϯ SE) in mean arterial pressure (MAP; 50 Ϯ 7 vs. 30 Ϯ 5 mmHg), popliteal blood velocity (15 Ϯ 3 vs. 5 Ϯ 1 cm/s), popliteal venous PCO2 (15 Ϯ 3 vs. 3 Ϯ 1 mmHg), and a greater decrease in popliteal venous pH (0.07 Ϯ 0.01 vs. 0.03 Ϯ 0.01), suggesting greater metabolic stimulation during dynamic contraction. Similarly, when peak tension was held constant and time was allowed to vary, dynamic contraction evoked a greater increase in blood velocity (13 Ϯ 1 vs. Ϫ1 Ϯ 1 cm/s) without causing any differences in other variables. To investigate the reflex contribution of mechanoreceptors, we stretched the hindlimb dynamically and statically at the same TTI. A larger reflex increase in MAP during dynamic stretch (32 Ϯ 8 vs. 24 Ϯ 6 mmHg) was observed when time was held constant, indicating greater mechanoreceptor stimulation. However, when peak tension was held constant, there were no differences in the reflex cardiovascular response to static and dynamic stretch. In conclusion, at comparable TTI, when peak tension is variable, dynamic muscle contraction causes larger cardiovascular responses than static contraction because of greater chemical and mechanical stimulation. However, when peak tensions are equivalent, static and dynamic contraction or stretch produce similar cardiovascular responses. static and dynamic muscle stretch; skeletal muscle blood flow; exercise; kao stimulator

INDIVIDUALS WITH CARDIOVASCULAR disease often are directed away from exercise involving static contraction, because it is thought that there is a greater reflex autonomic response associated with a larger afterload placed on the heart compared with dynamic contraction. Several attempts have been made to compare the cardiovascular response to both static and dynamic

contraction (1–3); however, comparisons between static and dynamic contraction have not been adequately investigated with the same muscle group performing equivalent amounts of work. Both static and rhythmic contraction of hindlimb muscles in anesthetized animals induce reflex increases in cardiac output, blood pressure, and heart rate (HR) (5, 6, 10, 12, 13). The magnitude of this reflex response is dependent on the amount of muscle mass and/or tension development, such that the greater the muscle mass involved or tension developed, the greater the pressor reflex (3, 5, 12). Previous studies suggest that static contraction causes greater pressor responses than those caused by dynamic contraction (9, 15, 19). This difference is reportedly due to continuous restriction of muscle blood flow during static contraction that results in greater accumulation of local metabolites and greater activation of chemosensitive muscle afferents (9, 15). However, differences in the energy of activation between these two types of contraction may play an important role because they can lead to differences in muscle metabolism and in the production and accumulation of local metabolites (4, 7, 17). Differential stimulation of muscle mechanoreceptors (intermittent vs. continuous) also may contribute to differences in the pressor response to dynamic contraction. Unfortunately, separating the effects of mechanoreceptors from those of metaboreceptors on the exercise pressor reflex can be difficult. Nevertheless, comparing the reflex cardiovascular response to passive dynamic and static muscle stretch provides a viable alternative because stretch is not associated with any metabolic changes (18). Although previous attempts have been made to compare cardiovascular responses with dynamic and static contractions (9, 15, 19), such comparisons have not been performed at equivalent workloads. Because the magnitude of the reflex cardiovascular response is directly related to the force production (15), adequate comparison of the response patterns during dynamic

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Samples were taken immediately before and after contraction or muscle stretch. Two sets of comparisons were made: one between the baseline values. Davis. The nerve was kept moist with mineral oil. Protocols Static versus dynamic contraction (TTI matched with time constant and variable peak tension). Surgical Preparation Adult cats of either sex (2. In six cats. 1). Static versus dynamic contraction/stretch (TTI matched with variable time and constant peak tension). and a catheter was directed in a retrograde fashion into the popliteal vein so that its tip was positioned just proximal to the triceps surae muscle group. whereas arterial pH was maintained by administering intravenous sodium bicarbonate (8. Each animal was placed in a spinal unit (David Kopf). Additional doses of ␣-chloralose (10 mg/kg) were administered as needed throughout the experiment. which was determined by integrating the area under the mean tension curves during the 30-s contraction period.4 ml/sample) to be withdrawn from the venous effluent of the triceps surae muscle group.4%) as needed.05. in part.35– 7. and respiration was maintained by a mechanical ventilator. PO2 Ͼ90 Torr.6 ml/cat) was obtained in each protocol. Fig. static contraction induces a greater reflex-pressor response than dynamic contraction. HR was assessed with a cardiotachometer (Gould 13–4515–65) that was triggered by the arterial blood pressure signal. Mean tension was displayed simultaneously through a lowpass filter (0. The order of contractions was randomized. 2). 100% oxygen was supplemented to maintain arterial PO2 Ͼ90 Torr. A total of four blood samples (1. even though popliteal blood velocity was significantly greater for dynamic than for static contraction. appears to be a reasonable quantitative technique for equating force produced during static and dynamic contraction. dynamic contractions were evoked to generate greater peak tension (5 Ϯ 1 vs. termed tensiontime index (TTI). the cats were paralyzed with vecuronium (3–5 mg/kg iv) at the end of the experiment. we stretched the hindlimb dynamically (1 Hz) and statically for 30 s to equivalent TTI (n ϭ 5 cats).3–5. The triceps surae was also contracted dynamically for 30 s by electrically stimulating the sciatic nerve with the modified Kao stimulator (60-Hz carrier frequency. we tested the following hypotheses: 1) when developed tension over time (TTI) is similar.35 Hz).025 ms). Average developed tension was equivalent to that produced by electrically induced contractions using the same stimulation parameters as described previously. 2) the smaller response to dynamic contraction is due. Statistical significance was accepted at PՅ0. In some animals. pH 7. METHODS right hindlimb was clamped in a fixed position. Tension produced by the contraction of the triceps surae was recorded as an index of overall hindlimb contraction. which tends to “wash out” local metabolites that induce the exercise pressor reflex. to greater skeletal muscle blood flow. If necessary.. This approach provides an accurate means to compare the cardiovascular response during these two disparate types of contraction within the same groups of muscle (1. RESULTS This study was approved by the Animal Use and Care Administrative Advisory Committee at the University of California. and the Achilles tendon was detached at the calcaneus bone and attached to an isometric force transducer (Grass FT-10). Therefore. the popliteal artery was dissected free of the surrounding tissue. Tension was expressed as average developed tension or TTI. Blood velocity was measured by a Doppler flow velocity meter (Triton Instruments. Dynamic contraction also induced significantly greater changes in venous effluent PCO2 and pH (Table 1).0 kg) were anesthetized with ketamine (25–30 mg/kg im) followed by ␣-chloralose (60–80 mg/kg iv).5 and 2. A catheter was placed in the left femoral vein for administration of drugs or fluids and in the left femoral artery for sampling arterial blood gases and for measurement of systemic arterial blood pressure. 2 Ϯ 1 kg. . Statistical analysis. and an electrode was attached to either a stimulus-isolation unit (Grass PSIU6) and a squarewave stimulator (Grass S88) for static contraction or a Kao. Static and dynamic contractions were evoked with voltages that varied between 1. This set of experiments was carried out to determine whether our observed differences in reflex responses were the result of differences in peak tension. and the right sciatic nerve was dissected free of surrounding tissue and placed on a shielded electrode. The trachea was intubated. To determine the role of mechanoreceptors in the reflex cardiovascular responses to both static and dynamic contraction. and a Doppler flow transducer was placed around the artery for assessment of blood velocity to the triceps surae.R1850 STATIC VS. 15). n ϭ 5 cats) during which peak tension and TTI were matched. To ensure that group III and IV muscle afferents were not stimulated.45. The Static Versus Dynamic Contraction TTI matched with time constant and variable peak tension. We conducted both static (30 s) and dynamic (60 s) contractions (n ϭ 6 cats) in addition to static (30 s) and dynamic stretch (60 s.e. modulated sinusoidally at 1 Hz). In seven cats. in seven cats.5 times the motor threshold. the length attained at full dorsiflexion). the triceps surae muscle was stretched to ϳ90–110% of its maximal in vivo muscle length (i. Arterial blood pressure was measured with a pressure transducer (Statham P23ID) attached to the arterial catheter. model 100) and was expressed in centimeters per second. All values are expressed as means Ϯ SE. This allowed blood samples (0. the triceps surae was contracted statically for 30 s by electrically stimulating the sciatic nerve at a frequency of 40 Hz (pulse duration ϭ 0. To match TTI between these two contractions over the same time interval (30 s). and the other between the peak changes in response to dynamic and static contraction or static and dynamic stretch using the Student’s paired t-test. On the basis of data from a previous study (18). sinusoidal wave stimulator for dynamic contractions. DYNAMIC CONTRACTION AT EQUIVALENT WORKLOADS and static exercise cannot be made accurately unless force production is similar. the saphenous vein was cannulated. Static versus dynamic stretch (TTI matched with time constant and variable peak tension). The integration of force over time. and the stimulus was repeated to demonstrate a lack of change in blood pressure or HR. Dynamic contraction evoked a greater increase in mean arterial pressure (MAP) compared with static contraction (Fig. Arterial blood gases and pH were maintained within the following ranges: PCO2 25–35 Torr.

In addition. greater reflex increases in MAP and HR were observed during dynamic compared with during static stretch (Fig. Static Versus Dynamic Stretch TTI matched with time constant and variable peak tension.STATIC VS. and mean tension. there were no differences in HR. In this protocol (n ϭ 6). pH. Dynamic contraction evoked increases in MAP that were similar to those caused by static contraction (Fig. Beats/ min. bpm.0 Ϯ 0. absolute tension.5 kg) contraction. During 30 s of passive stretch (n ϭ 5). heart rate.0 Ϯ 0. 4). No significant differences were observed in venous lactate concentration ([lactate]) or PO2. or [lactate] (Table 1). despite greater increases in popliteal blood velocity during dynamic contraction. TTI matched with variable time and constant peak tension. 3). 1). peak developed tension was matched during static (2. DYNAMIC CONTRACTION AT EQUIVALENT WORKLOADS R1851 Fig. Variables shown are mean arterial pressure (MAP). but the time of contraction was varied (30 s static vs. PO2. 1. 60 s dynamic) to equate TTI (Fig. PCO2. Original record of the cardiovascular response to static and dynamic contraction at matched tension-time index with peak tension held constant (A) and with time held constant (B).5 kg) and dynamic (2. .

There were no significant differences in the reflex cardiovascular response to static and dynamic stretch during these conditions (Fig.R1852 STATIC VS.32 Ϯ 0. Thus the greater cost of muscle activation and higher peak tensions during dynamic contraction probably augmented the production of muscle metabolites.4* 0.9 Ϯ 0. The results of our study did not confirm our hypothesis. However.2 37 Ϯ 4 66 Ϯ 5* 29 Ϯ 4 27 Ϯ 1 36 Ϯ 9 9Ϯ4 7. DYNAMIC CONTRACTION AT EQUIVALENT WORKLOADS DISCUSSION Fig. venous PCO2 and Hϩ levels during dynamic contraction were actually higher than during static contraction. Peak changes (⌬. . TTI matched with variable time and constant peak tension.01† 1.01 Values are means Ϯ SE. Change.0 Ϯ 0.02 7.30 Ϯ 0. dynamic.vs.01 7.0 Ϯ 0. 7.02*† Ϫ0.1 Ϯ 0. heart rate.31 Ϯ 0.2 35 Ϯ 2 60 Ϯ 6* 25 Ϯ 3 28 Ϯ 2 37 Ϯ 2 9Ϯ1 7. 17. of cats. * P Ͻ 0.6 Ϯ 0. and variable peak tension. mM Precontraction Postcontraction ⌬ PCO2. n.4 2. When TTI was matched by maintaining similar peak tensions and increasing contraction time during dy- Table 1. Numbers below histograms represent baseline values.02 Ϫ0.13 Ϯ 0. greater stimulation of metaboreceptors most likely contributed to the larger reflex-pressor response to dynamic contraction.01 1. Instead. we found that the pressor response to dynamic contraction was not reduced compared with static contraction under these conditions. Overall.20 Ϯ 0.3 2. No significant change in popliteal artery blood velocity was observed (Fig.4 Ϯ 0.1 41 Ϯ 4 44 Ϯ 3 3Ϯ1 34 Ϯ 5 34 Ϯ 4 0Ϯ1 7.6 Ϯ 0.2 Ϯ 0. Metabolic responses to dynamic and static contraction at equivalent tension-time index Time Constant (n ϭ 7) Static Dynamic Peak Tension Constant (n ϭ 6) Static Dynamic Lactate. mmHg Precontraction Postcontraction ⌬ PO2. the higher muscle blood flow associated with the greater peak tension likely limited accumulation of these local metabolites. by holding contraction time constant and by increasing peak tension during dynamic contraction to equate TTI.8 Ϯ 0. *P Ͻ 0.6 Ϯ 0. static vs.28 Ϯ 0. intermittent contractions have previously been shown to cause greater increases in energy metabolism and muscle metabolite production than static contraction (4.1 40 Ϯ 3 55 Ϯ 4*† 15 Ϯ 3† 34 Ϯ 5 31 Ϯ 3 Ϫ3 Ϯ 2 7.25 Ϯ 0. This difference in the cardiovascular response occurred despite the fact that active muscle blood velocity was significantly higher during dynamic contraction. constant time (30 s). However.1* 0. postcontraction. dynamic.4* 0.05. pre. 2.02 7. 5). ⌬. Moreover.3 1.1* 0. mmHg Precontraction Postcontraction ⌬ pH Precontraction Postcontraction ⌬ 1.01* Ϫ0.4 2.10 Ϯ 0.07 Ϯ 0.31 Ϯ 0.01 1. These differences appear to be related to greater ATP requirements for the initial activation of the muscle during intermittent contraction (7). 18). We had proposed that static contraction would elicit a greater reflex-pressor response than dynamic contraction at comparable workloads.03 Ϯ 0.02 7. Short duration. We expected that the greater blood flow during dynamic contraction would result in less activation of the exercise-pressor reflex both because of a greater supply of oxygen and because of greater “metabolite washout” in the active muscle.01* Ϫ0.05.7 Ϯ 0. the pressor response to dynamic contraction was greater than that caused by static contraction. static vs. and mean blood velocity during static and dynamic contraction (n ϭ 7) at equivalent tension-time indexes. means Ϯ SE) in MAP. † P Ͻ 0. 4). no.05.2 Ϯ 0.18 Ϯ 0.

*P Ͻ 0. In all likelihood. this technique primarily stimulated group III muscle afferent nerve endings. at comparable TTI. because these afferents are believed to be primarily mechanically sensitive.05. Greater metabolite production during dynamic contraction presumably was due to the greater metabolic cost of repetitive muscle activation. dynamic. heart rate. 4. static vs. . *P Ͻ 0. there are both a greater production and removal of metabolites during dynamic contraction that result in no net accumulation compared with static contraction. revealed no differences between the two modes of stretch. Although dis- Fig. However. myocardial contractility. However. namic contraction. and HR. static vs. Fig. 3. 60 s dynamic).STATIC VS. a greater pressor response occurred during dynamic stretch. dynamic. Comparison of the pressor responses induced by static and dynamic stretch at the same TTI. However. Peak changes (means Ϯ SE) in MAP. when a greater peak tension was produced so that time could be held constant during dynamic and static stretch. and variable peak tension. constant peak tension. when peak tension was matched. and mean blood velocity during static and dynamic contraction (n ϭ 6) at equivalent tension-time indexes. this technique allowed us to isolate the effects of mechanoreceptor activation from those caused by metaboreceptor activation.05. we observed similar increases in blood pressure and venous metabolite concentrations during both types of contraction. because there is greater activation of metaboreceptors. and mean blood velocity during static and dynamic stretch (n ϭ 5) at equivalent tension-time indexes. differential activation of mechanoreceptors could also be involved. heart rate. DYNAMIC CONTRACTION AT EQUIVALENT WORKLOADS R1853 Because passive stretch of skeletal muscle reflexly increases blood pressure. blood velocity was still considerably greater during dynamic contraction. These findings also provide support for the contention that dynamic contraction causes a greater pressor response than static contraction (when TTI is equated by holding time constant). whereas group IV muscle afferents are considered to be predominately chemically sensitive (10). Nevertheless. We examined this possibility by comparing dynamic and static stretch using the same protocols in which dynamic and static contractions were compared. These patterns of response were consistent with those observed during static and dynamic contraction under similar conditions. Numbers below histograms represent baseline values. without changing muscle metabolite concentrations (18). constant time (30 s). Numbers below histograms represent baseline values. Peak changes (means Ϯ SE) in MAP. and variable time (30 s static vs. increased accumulation of these substances appears to have been offset by higher blood flows. These findings support the premise that when peak tension is equal.

may not accurately reflect the level of activation of the sympathetic nervous system (16). Thus sequential activation of muscle mechanoreceptors by the Kao stimulator may influence the magnitude of the corresponding pressor response compared with contraction evoked by unmodified stimulation in which peak intensity is reached instantaneously. For example. charge activity of mechanoreceptors can be altered by the frequency of muscle contraction in that continuous activation may lead to adaptation or decreased firing (9). therefore. oxygen uptake was used to assess work intensity. especially if there are large differences in the amount of active muscle mass involved. because these changes in autonomic activity are more dependent on local metabolic changes in exercising muscle rather than whole body metabolism. a level of static exercise (isometric leg extension) that requires less oxygen consumption than dynamic exercise (cycle ergometery) can induce greater increases in muscle sympathetic nerve activity compared with dynamic contraction (16). 60 s dynamic). The problem with this approach is that whole body metabolism. Therefore. and variable time (30 s static vs. This pattern of activation is similar to that present during voluntary contraction and causes a pattern of mechanoreceptor activation that is different from that induced with a standard square-wave stimulator. heart rate. because muscle tension usually begins to decline within ϳ15–20 s. Most likely. our data suggest that the smaller pressor response to static contraction when time was held constant was not due to adaptation. who showed that intermittent tetanic contraction evokes synchronized renal nerve discharge. and mean blood velocity during static and dynamic stretch (n ϭ 5) at equivalent tension-time indexes. Previous studies in humans have reported comparable pressor responses to static and dynamic contraction (2. That is why a contraction period of only 30 s was selected. 3). 5. comparison of the cardiovascular responses during these two forms of exercise at the same level of oxygen consumption could lead to inaccurate conclusions. It also should be noted that cardiovascular response to electrically induced muscle contraction does reach a true steady state. because it allowed us to more closely simulate rhythmic contractions as they occur naturally (8). Additional studies employing afferent unit recording will be required to confirm this potential differential activation of muscle mechanosensitive receptors during dynamic muscular contraction. DYNAMIC CONTRACTION AT EQUIVALENT WORKLOADS Fig. whole body metabolism (oxygen uptake) does not correlate closely with sympathetic nerve activity during exercise. the pressor response to contraction usually peaks within 20–45 s and does not achieve a plateau for Ͼ5–10 s. . Therefore. when peak tension was varied. because the stimulus intensity increases and decreases gradually. (20). This outcome is particularly true for static contraction (see Fig.R1854 STATIC VS. The Kao stimulator produces a sinusoidal stimulus that increases slowly from low to peak intensity and then gradually decreases in a similar fashion (8). This potential contribution of mechanoreceptors is in agreement with Victor et al.and mechanoreceptors contribute to the larger reflex-pressor response during dynamic contraction at comparable workloads when peak tension is allowed to vary. However. We have found that even when the decline in tension is small. as indicated by oxygen uptake. 1). supporting a role for mechanoreceptor afferents in the reflex cardiovascular response to dynamic contraction. we used a modified Kao stimulator (see METHODS) to evoke dynamic contractions. To evoke dynamic contraction in this study. greater mechanoreceptor stimulation may have occurred due to higher peak tension development during dynamic contraction such that a greater reflex sympathetic nerve activation and a greater pressor response occurred. we believe that our findings indicate that greater activation of both metabo. Numbers below histograms represent baseline values. constant peak tension. workloads were equated by matching oxygen uptake. because there was no difference between the cardiovascular response to static and dynamic contraction when peak tension was held constant. In these studies. Peak changes (means Ϯ SE) in MAP.

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