A comparison of the triceps surae and residual muscle moments at the ankle during cycling

The rigid linked system model and principles of inverse dynamics have been widely used to calculate residual muscle moments during various activities. EMG driven models and optimization algorithms have also been presented in the literature in efforts to estimate skeletal muscle forces and evaluate their possible contribution to the residual muscle moment. Additionally, skeletal muscle-tendon forces have been measured, directly, in both animals and humans. The purpose of this investigation was to calculate the moment produced by the triceps surae muscles and compare it to the residual muscle moment at the ankle during cycling at three power outputs (90, 180 and 270 W). Inferences were made regarding the potential contribution made by each triceps surae component to the tendon force using EMG and muscle-tendon length changes. A buckle-type transducer was surgically implanted on the right Achilles tendon of one male subject. Achilles tendon forces measured in vivo were multiplied by their corresponding moment arms to yield the triceps surae moment during the three working conditions. Moment arm lengths were obtained in a separate experiment using magnetic resonance imaging (MRI). Pedal reaction forces, body segment accelerations (determined from high speed film), and appropriate mass parameters served as input to the inverse solution. The triceps surae moment was temporally in phase with and consistently represented approximately 65% of the residual muscle moment at the ankle. These data demonstrate the feasibility of using implanted transducers in human subjects and provide a greater understanding of musculoskeletal mechanics during normal human movements.     

Comparison of the exercise pressor reflex between forelimb and hindlimb muscles in cats

In thirteen cats anesthetized with alpha-chloralose, we compared the cardiovascular and ventilatory responses to both static contraction and tendon stretch of a hindlimb muscle group, the triceps surae, with those to contraction and stretch of a forelimb muscle group, the triceps brachii. Static contraction and stretch of both muscle groups increased mean arterial pressure and heart rate, and the responses were directly proportional to the developed tension. The cardiovascular increases, however, were significantly greater (P < 0.05) when the triceps brachii muscles were contracted or stretched than when the triceps surae muscles were contracted or stretched, even when the tension developed by either maneuver was corrected for muscle weight. Likewise, the ventilatory increases were greater when the triceps brachii muscles were stretched than when the triceps surae muscles were stretched. Contraction of either muscle group did not increase ventilation. Our results suggest that in the anesthetized cat the cardiovascular responses to both static contraction and tendon stretch are greater when arising from forelimb muscles than from hindlimb muscles.     

Soleus aponeurosis strain distribution following chronic unloading in humans: an in vivo MR phase-contrast study.

The in vivo strain properties of human skeletal muscle-tendon complexes are poorly understood, particularly following chronic periods of reduced load bearing. We studied eight healthy volunteers who underwent 4 wk of unilateral lower limb suspension (ULLS) to induce chronic unloading. Before and after the ULLS, maximum isometric ankle plantar flexion torque was determined by using a magnetic resonance (MR)-compatible dynamometry. Volumes of the triceps surae muscles and strain distribution of the soleus aponeurosis and the Achilles tendon at a constant submaximal plantar flexion (20% pre-maximal voluntary contraction) were measured by using MRI and velocity-encoded, phase-contrast MRI techniques. Following ULLS, volumes of the soleus and the medial gastrocnemius and the maximum isometric ankle plantar flexion (maximum voluntary contraction) decreased by 5.5+/-1.9, 7.5+/-2.7, and 48.1+/-6.1%, respectively. The strain of the aponeurosis along the length of the muscle before the ULLS was 0.3+/-0.3%, ranging from -1.5 to 2.7% in different locations of the aponeurosis. Following ULLS, the mean strain was -6.4+/-0.3%, ranging from -1.6 to 1.3%. The strain distribution of the midregion of the aponeurosis was significantly influenced by the ULLS, whereas the more distal component showed no consistent changes. Achilles tendon strain was not affected by the ULLS. These results raise the issue as to whether these changes in strain distribution affect the functional properties of the triceps surae and whether the probability of strain injuries within the triceps surae increases following chronic unloading in those regions of this muscle complex in which unusual strains occur.     

Medial gastrocnemius muscle fascicle active torque-length and Achilles tendon properties in young adults with spastic cerebral palsy

Individuals with spastic cerebral palsy (CP) typically experience muscle weakness. The mechanisms responsible for muscle weakness in spastic CP are complex and may be influenced by the intrinsic mechanical properties of the muscle and tendon. The purpose of this study was to investigate the medial gastrocnemius (MG) muscle fascicle active torque-length and Achilles tendon properties in young adults with spastic CP. Nine relatively high functioning young adults with spastic CP (GMFCS I, 17±2 years) and 10 typically developing individuals (18±2 years) participated in the study. Active MG torque-length and Achilles tendon properties were assessed under controlled conditions on a dynamometer. EMG was recorded from leg muscles and ultrasound was used to measure MG fascicle length and Achilles tendon length during maximal isometric contractions at five ankle angles throughout the available range of motion and during passive rotations imposed by the dynamometer. Compared to the typically developing group, the spastic CP group had 33% lower active ankle plantarflexion torque across the available range of ankle joint motion, partially explained by 37% smaller MG muscle and 4% greater antagonistic co-contraction. The Achilles tendon slack length was also 10% longer in the spastic CP group. This study confirms young adults with mild spastic CP have altered muscle–tendon mechanical properties. The adaptation of a longer Achilles tendon may facilitate a greater storage and recovery of elastic energy and partially compensate for decreased force and work production by the small muscles of the triceps surae during activities such as locomotion.     

Three-Dimensional Ankle Moments and Nonlinear Summation of Rat Triceps Surae Muscles

The Achilles tendon and epimuscular connective tissues mechanically link the triceps surae muscles. These pathways may cause joint moments exerted by each muscle individually not to sum linearly, both in magnitude and direction. The aims were (i) to assess effects of sagittal plane ankle angle (varied between 150° and 70°) on isometric ankle moments, in both magnitude and direction, exerted by active rat triceps surae muscles, (ii) to assess ankle moment summation between those muscles for a range of ankle angles and (iii) to assess effects of sagittal plane ankle angle and muscle activation on Achilles tendon length. At each ankle angle, soleus (SO) and gastrocnemius (GA) muscles were first excited separately to assess ankle-angle moment characteristics and subsequently both muscles were excited simultaneously to investigate moment summation. The magnitude of ankle moment exerted by SO and GA, the SO direction in the transverse and sagittal planes, and the GA direction in the transverse plane were significantly affected by ankle angle. SO moment direction in the frontal and sagittal planes were significantly different from that of GA. Nonlinear magnitude summation varied between 0.6±2.9% and −3.6±2.9%, while the nonlinear direction summation varied between 0.3±0.4° and −0.4±0.7° in the transverse plane, between 0.5±0.4° and 0.1±0.4° in the frontal plane, and between 3.0±7.9° and 0.3±2.3° in the sagittal plane. Changes in tendon length caused by SO contraction were significantly lower than those during contraction of GA and GA+SO simultaneously. Thus, moments exerted by GA and SO sum nonlinearly both in the magnitude and direction. The limited degree of nonlinear summation may be explained by different mechanisms acting in opposite directions.     

Effect of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit

Findings from animal experiments are sometimes contradictory to the idea that the tendon structure is a simple elastic spring in series with muscle fibers, and suggest influence of muscle contraction on the tendon mechanical properties. The purpose of the present study was to investigate the influence of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit. For seven subjects, ankle dorsiflexion was performed without (passive condition) and with contraction of plantar flexor muscles (eccentric conditions, at 3 contraction levels) on an isokinetic dynamometer. Deformation of the Achilles tendon during each trial was measured using ultrasonography. The Achilles tendon force corresponding to the tendon elongation of 10mm in the passive condition was significantly smaller than those in the eccentric conditions (p<0.05 or p<0.01). Within the eccentric conditions, the Achilles tendon force corresponding to the tendon elongation of 10mm was significantly greater in the maximal contraction level than those in submaximal eccentric conditions (p<0.05 or p<0.01). In addition, the tendon stiffness was greater in higher contraction levels (p<0.05 or p<0.01). Present results suggest that the human tendon structure is not a simple elastic spring in series with muscle fibers.     

A 3D model of the Achilles tendon to determine the mechanisms underlying nonuniform tendon displacements

The Achilles is the thickest tendon in the body and is the primary elastic energy-storing component during running. The form and function of the human Achilles is complex: twisted structure, intratendinous interactions, and differential motor control from the triceps surae muscles make Achilles behavior difficult to intuit. Recent in vivo imaging of the Achilles has revealed nonuniform displacement patterns that are not fully understood and may result from complex architecture and musculotendon interactions. In order to understand which features of the Achilles tendon give rise to the nonuniform deformations observed in vivo, we used computational modeling to predict the mechanical contributions from different features of the tendon. The aims of this study are to: (i) build a novel computational model of the Achilles tendon based on ultrashort echo time MRI, (ii) compare simulated displacements with published in vivo ultrasound measures of displacement, and (iii) use the model to elucidate the effects of tendon twisting, intratendon sliding, retrocalcaneal insertion, and differential muscle forces on tendon deformation. Intratendon sliding and differential muscle forces were found to be the largest factors contributing to displacement nonuniformity between tendon regions. Elimination of intratendon sliding or muscle forces reduced displacement nonuniformity by 96% and 85%, respectively, while elimination of tendon twist and the retrocalcaneal insertion reduced displacement nonuniformity by only 35% and 3%. These results suggest that changes in the complex internal structure of the tendon alter the interaction between muscle forces and tendon behavior and therefore may have important implications on muscle function during movement.     

Triceps surae activation is altered in male runners with Achilles tendinopathy

Achilles tendinopathy is a common injury in running sports however the exact etiology of Achilles injury is still unclear. In recent years, altered neuromotor recruitment patterns of the triceps surae have been hypothesized to create differential intra-tendinous loads leading to pathology; however, this hypothesis has not been investigated. Further, the effect foot orthoses may have on neuromotor recruitment of the triceps surae in Achilles tendinopathy has not been investigated. Methods: The electromyographic activity of the triceps surae was recorded during an over-ground running task. Fifteen Achilles injured participants and 19 asymptomatic controls were assessed in a footwear only condition. The Achilles injured participants were also assessed running in a pre-fabricated foot orthoses. Results: In Achilles injured participants, there was a significant difference between soleus and lateral gastrocnemius offset times during running compared to the asymptomatic controls (p < 0.05). There were no significant differences in triceps surae muscle activity between the footwear only and footwear and orthoses condition in the Achilles injured participants. Conclusions: The finding that triceps surae activity is altered in participants with Achilles tendinopathy may have clinical importance as it suggests that intra-tendinous loads are altered which may contribute to pathological changes. Further, foot orthoses have no immediate effect on the neuromotor control of the triceps surae.     

Ethnic differences in triceps surae muscle-tendon complex and walking economy

The purposes of this study were to (a) determine whether structural differences in triceps surae muscle-tendon complex and walking economy exist between 14 African American and 19 Caucasian sedentary women and (b) determine whether muscle-tendon parameters are associated with walking economy. African American and Caucasian subjects were matched on body weight, height, and body composition. Muscle-tendon parameters were determined by magnetic resonance imaging and walking economy was evaluated at 4.8 km.h(-1). Medial gastrocnemius and total triceps surae muscle shape were different across ethnicity despite no ethnic differences in plantar flexion strength or in maximal cross-sectional area for any triceps surae muscles. African American women had shorter gastrocnemius muscles and longer tendons and performed walking more economically. Tendon length was the only variable related to walking economy. No ethnic differences were observed in walking economy after adjusting for tendon length. Data show gastrocnemius tendon length is related to level walking and longer gastrocnemius tendons may partly explain more economical walking in African American women. These preliminary findings indicate the structure of the muscle-tendon complex could be a factor partially accounting for reported ethnic differences in certain types of athletic-related performance.     

Aging effects on the Achilles tendon moment arm during walking

The Achilles tendon (AT) moment arm transforms triceps surae muscle forces into a moment about the ankle which is critical for functional activities like walking. Moreover, the AT moment arm changes continuously during walking, as it depends on both ankle joint rotation and triceps surae muscle loading (presumably due to bulging of the muscle belly). Here, we posit that aging negatively effects the architecturally complex AT moment arm during walking, which thereby contributes to well-documented reductions in ankle moment generation during push-off. We used motion capture-guided ultrasound imaging to quantify instantaneous variations in the AT moment arms of young (23.9 ± 4.3 years) and older (69.9 ± 2.6 years) adults during walking, their dependence on triceps surae muscle loading, and their association with ankle moment generation during push-off. Older adults walked with 11% smaller AT moment arms and 11% smaller peak ankle moments during push-off than young adults. Moreover, as hypothesized, these unfavourable changes were significantly and positively correlated (r² = 0.38, p < 0.01). More surprisingly, aging attenuated load-dependent increases in the AT moment arm (i.e., those between heel-strike and push-off at the same ankle angle); only young adults exhibited a significant increase in their AT moment arm due to triceps surae muscle-loading. Age-associated reductions in triceps surae volume or activation, and thus muscle bulging during force generation, may compromise the mechanical advantage of the AT during the critical push-off phase of walking in older adults. Thus, strategies to restore and/or improve locomotor performance in our aging population should consider these functionally important changes in musculoskeletal behavior.     

Variation in the human Achilles tendon moment arm during walking

The Achilles tendon (AT) moment arm is an important determinant of ankle moment and power generation during locomotion. Load and depth-dependent variations in the AT moment arm are generally not considered, but may be relevant given the complex triceps surae architecture. We coupled motion analysis and ultrasound imaging to characterize AT moment arms during walking in 10 subjects. Muscle loading during push-off amplified the AT moment arm by 10% relative to heel strike. AT moment arms also varied by 14% over the tendon thickness. In walking, AT moment arms are not strictly dependent on kinematics, but exhibit important load and spatial dependencies.     

Disuse atrophy increases the muscle mechanoreflex in rats.

We investigated the effect of disuse atrophy on the magnitude of the muscle mechanoreflex. The left leg of eight rats (6-7 wk, male) was put in a plaster cast for 1 wk. The rats were decerebrated at the midcollicular level. We recorded the pressor and cardioaccelerator responses to 30-s stretch of the calcaneal tendon, which selectively stimulated the muscle mechanosensitive receptors in the left atrophied and right control triceps surae muscles. Atrophied muscles showed significantly lower mass control muscles (1.0 +/- 0.1 vs. 1.4 +/- 0.1 g; P < 0.05). At the same stretch tension (229 +/- 20 g), the pressor response to stretch was significantly greater in the atrophied muscles than in the control muscles (13 +/- 3 vs. 4 +/- 2 mmHg, P < 0.05). The cardioaccelerator response was not significantly different (8 +/- 4 vs. 4 +/- 2 beats/min). Comparing responses at the same relative tension (57 +/- 6 vs. 51 +/- 8% of maximal tension), the pressor response was still significantly greater in the atrophied triceps surae than in the control (14 +/- 4 vs. 4 +/- 2 mmHg; P < 0.05). These results suggest that disuse atrophy increases the magnitude of muscle mechanoreflex.     

Effect of habitual running on human Achilles tendon load-deformation properties and cross-sectional area.

Whether the cross-sectional area (CSA) and mechanical properties of the human Achilles tendon change in response to habitual exercise remains largely unexplored. The present study evaluated the CSA and contraction-induced displacement of the aponeurosis-tendon complex of the triceps surae in 11 untrained subjects before (tests 1 and 2) and after (test 3) approximately 9 mo of regular running ( approximately 78 training sessions). Displacement of the tendon-aponeurosis complex obtained by ultrasonography; electromyography of the gastrocnemius, soleus, and dorsiflexor muscles; and joint angular rotation were recorded during graded isometric plantarflexion ramps. Tendon CSA and moment arm were measured by using MRI, and tendon force was calculated from joint moments and tendon moment arm. A treadmill test was used to determine submaximal oxygen consumption (Vo2) at a given speed and maximal Vo2. The total running duration was approximately 43 h, distributed over 34 wk. Maximal Vo2 increased 8.6% (P < 0.01), and submaximal Vo2 decreased 6.2% (P < 0.05). Tendon-aponeurosis displacement during maximal voluntary contraction was unchanged (tests 1-3, 5.2 +/- 0.6, 5.2 +/- 0.5, and 5.3 +/- 0.4 mm, respectively) and yielded a structural stiffness of 365 +/- 50, 358 +/- 40, and 384 +/- 52 N/mm for tests 1-3, respectively (P > 0.05). Tendon CSA also remained unchanged (tests 1-3, 34.2 +/- 2.2, 33.9 +/- 2.2, and 33.8 +/- 2.1 mm2, respectively). In conclusion, a total training stimulus of approximately 9 mo of running in previously untrained subjects was adequate to induce significant cardiovascular improvements, although it did not result in any changes in the mechanical properties of the triceps surea tendon-aponeurosis complex or in the dimensions of Achilles tendon.     

Changes in stretch reflexes and muscle stiffness with age in prepubescent children.

Musculo-articular stiffness of the triceps surae (TS) increases with age in prepubescent children, under both passive and active conditions. This study investigates whether these changes in muscle stiffness influence the amplitude of the reflex response to muscle stretch. TS stiffness and reflex activities were measured in 46 children (7-11 yr old) and in 9 adults. The TS Hoffmann reflex (H reflex) and T reflex (tendon jerk) in response to taping the Achilles tendon were evaluated at rest and normalized to the maximal motor response (Mmax). Sinusoidal perturbations of passive or activated muscles were used to evoke stretch reflexes and to measure passive and active musculoarticular stiffness. The children's Hmax-to-Mmax ratio did not change with age and did not differ from adult values. The T-to-Mmax ratio increased with age but remained significantly lower than in adults. Passive stiffness also increased with age and was correlated with the T-to-Mmax ratio. Similarly, the children's stretch reflex and active musculoarticular stiffness were significantly correlated and increased with age. We conclude that prepubescent children have smaller T reflexes and stretch reflexes than adults, and the lower musculoarticular stiffness is mainly responsible for these smaller reflexes, as indicated by the parallel increases in reflex and stiffness.     

The role of the stretch reflex in the gastrocnemius muscle during human locomotion at various speeds.

In the present study, the fascicle length (L(fa)) of the human medial gastrocnemius (MG) muscle was monitored to evaluate possible input from the short-latency stretch reflex (SLR) during the stance phase of running and to examine its timing at various running speeds. Eight subjects ran at 2.0, 3.5, 5.0, and 6.5 m/s. The L(fa) was measured with the high-speed ultrasound fascicle scanning together with kinematics and myoelectrical activities. The amplitudes and onset latency of SLR activities were determined. During ground contact, the sudden MG fascicle stretch occurred during the early contact at all running speeds. This was followed by the fascicle shortening. The timing of fascicle stretch depended on running speed and type of foot contact. In slower speed conditions (2.0, 3.5, 5 m/s), the MG fascicle stretch and the corresponding SLR activities occurred during the middle of the braking phase. In fast-speed running (6.5 m/s), however, the MG fascicle stretch occurred later compared with the lower speed. The corresponding SLR activities occurred significantly later at the end of the braking phase. In addition to the clear demonstration of the different timings of SLR in MG during ground contact of running, the results imply that the role of the MG SLR during the stance phase of running can be different between fast- and slow-speed running conditions.     

短跑训练对运动员跟腱影响的超声测量研究

目的：研究短跑训练对运动员跟腱的影响。方法：选择从事运动训练4-6年的健康男、女短跑运动员为实验组（n=12）,同时选同龄健康非运动员男、女为对照组（n=12）,采用高频超声检测平静状态跟腱长度、横截面积,以及小腿三头肌等长收缩最大力量跖屈跟腱长度的变化。结果：跟腱长度：对照组男子168.5±9.2mm,女子162.4±9.8mm,实验组男子170.9±10.7mm,女子164.0±7.0mm。实验组和对照组组内、组间差别均无统计学意义。跟腱横截面积：对照组男子62.2±6.2mm2,女子47.1±4.5mm2,实验组男子65.6±2.9mm2,女子49.6±1.9mm2。同组内男子比女子跟腱横截面积大,差别有统计学意义（P＆lt;0.05）,但实验组和对照组组间差别无统计学意义。小腿三头肌等长收缩最大力跖屈时跟腱拉长值：对照组男子7.6±3.6mm,女子4.9±2.8mm,实验组男子11.1±2.9mm,女子7.9±3.1mm。男子比女子跟腱拉长值大、实验组比对照组拉长值大,差别均有统计学意义（P＆lt;0.05）。结论：短跑训练可增加跟腱可拉伸长度,跟腱的长度和横截面积未发生明显变化。高频超声可作为重要的测量手段用于...     

Estimates of Achilles tendon moment arm differ when axis of ankle rotation is derived from ankle motion

The plantarflexor moment arm of the Achilles tendon determines the mechanical advantage of the triceps surae and also indirectly affects muscle force generation by setting the amount of muscle-tendon shortening per unit of ankle joint rotation. The Achilles tendon moment arm may be determined geometrically from an axis (or center) of joint rotation and the line of action of the tendon force, but such moment arms may be sensitive to the location of the joint axis. Using motion analysis to track an ultrasound probe overlying the Achilles tendon along with markers on the shank and foot, we measured Achilles tendon moment arm during loaded and unloaded dynamic plantarflexion motions in 15 healthy subjects. Three representations of the axis or center of rotation of the ankle were considered: (1) a functional axis, defined by motions of the foot and shank; (2) a transmalleolar axis; and (3) a transmalleolar midpoint. Moment arms about the functional axis were larger than those found using the transmalleolar axis and transmalleolar midpoint (all p < 0.001). Moment arms computed with the functional axis increased with plantarflexion angle (all p < 0.001), and increased with loading in the most plantarflexed position (p < 0.001) but these patterns were not observed when either using a transmalleolar axis or transmalleolar midpoint. Functional axis moment arms were similar to those estimated previously using magnetic resonance imaging, suggesting that using a functional axis for ultrasound-based geometric estimates of Achilles tendon moment arm is an improvement over landmark-based methods.     

Muscle mechanosensitive receptors close to the myotendinous junction of the Achilles tendon elicit a pressor reflex.

Feedback regulation by activation of mechanosensitive afferents in the exercising muscle causes the cardiovascular and sympathetic nerve responses, which follow tension development and are almost identical between static contraction and passive stretch. The precise location of the mechanoreceptors contributing to the exercise pressor reflex, however, remained unknown. To test the hypothesis that the mechanoreceptors will be located around the myotendinous junction to monitor a change in muscle tension than a change in muscle length, we examined the reflex cardiovascular responses to passive stretch of the triceps surae muscle in anesthetized rats with three interventions; systemic injection of gadolinium, cutting the Achilles tendon, and local injection of lidocaine into the myotendinous junction. Gadolinium (42 micromol/kg iv) blunted the increases in heart rate and mean arterial blood pressure during passive stretch by 36 and 22-26%, respectively, suggesting that the reflex cardiovascular responses were evoked by stimulation of muscle mechanosensitive receptors. The cardiovascular responses to passive stretch were not different between the cut Achilles tendon and the intact tendon in the same rats, suggesting that any mechanoreceptors, terminated in the more distal part of the tendon, did not contribute to the reflex cardiovascular responses. Lidocaine (volume, 0.04-0.1 ml) injected into the myotendinous junction blunted the stretch-induced increases in heart rate and mean arterial blood pressure by 37-49 and 27-34%, respectively. We conclude that the muscle mechanosensitive receptors evoking the reflex cardiovascular responses at least partly locate at or close to the myotendinous junction of the Achilles tendon.     

Gadolinium attenuates exercise pressor reflex in cats

The exercise pressor reflex, which arises from the contraction-induced stimulation of group III and IV muscle afferents, is widely believed to be evoked by metabolic stimuli signaling a mismatch between blood/oxygen demand and supply in the working muscles. Nevertheless, mechanical stimuli may also play a role in evoking the exercise pressor reflex. To determine this role, we examined the effect of gadolinium, which blocks mechanosensitive channels, on the exercise pressor reflex in both decerebrate and alpha-chloralose-anesthetized cats. We found that gadolinium (10 mM; 1 ml) injected into the femoral artery significantly attenuated the reflex pressor responses to static contraction of the triceps surae muscles and to stretch of the calcaneal (Achilles) tendon. In contrast, gadolinium had no effect on the reflex pressor response to femoral arterial injection of capsaicin (5 microg). In addition, gadolinium significantly attenuated the responses of group III muscle afferents, many of which are mechanically sensitive, to both static contraction and to tendon stretch. Gadolinium, however, had no effect on the responses of group IV muscle afferents, many of which are metabolically sensitive, to either static contraction or to capsaicin injection. We conclude that mechanical stimuli arising in contracting skeletal muscles contribute to the elicitation of the exercise pressor reflex.     

Role played by purinergic receptors on muscle afferents in evoking the exercise pressor reflex.

The exercise pressor reflex is believed to be evoked, in part, by multiple metabolic stimuli that are generated when blood supply to exercising muscles is inadequate to meet metabolic demand. Recently, ATP, which is a P2 receptor agonist, has been suggested to be one of the metabolic stimuli evoking this reflex. We therefore tested the hypothesis that blockade of P2 receptors within contracting skeletal muscle attenuated the exercise pressor reflex in decerebrate cats. We found that popliteal arterial injection of pyridoxal phosphate-6-azophenyl-2',4'-disulfonic acid (PPADS; 10 mg/kg), a P2 receptor antagonist, attenuated the pressor response to static contraction of the triceps surae muscles. Specifically, the pressor response to contraction before PPADS averaged 36 +/- 3 mmHg, whereas afterward it averaged 14 +/- 3 mmHg (P < 0.001; n = 19). In addition, PPADS attenuated the pressor response to postcontraction circulatory occlusion (P < 0.01; n = 11). In contrast, popliteal arterial injection of CGS-15943 (250 micro g/kg), a P1 receptor antagonist, had no effect on the pressor response to static contraction of the triceps surae muscles. In addition, popliteal arterial injection of PPADS but not CGS-15943 attenuated the pressor response to stretch of the calcaneal (Achilles) tendon. We conclude that P2 receptors on the endings of thin fiber muscle afferents play a role in evoking both the metabolic and mechanoreceptor components of the exercise pressor reflex.