Mechanical advantage of sternomastoid and scalene muscles in dogs

Legrand, Alexandre, Vincent Ninane, and André DeTroyer. Mechanical advantage of sternomastoid and scalene muscles in dogs. J. Appl. Physiol. 82(5):1517-1522, 1997.Theoretical studies have led to the predictionthat the maximal effect of a given respiratory muscle on airway openingpressure (Pao) is the product of muscle mass, the maximal active muscletension per unit cross-sectional area, and the fractional change inmuscle length per unit volume increase of the relaxed chest wall. It has previously been shown that the parasternal intercostals behave inagreement with this prediction (A. De Troyer, A. Legrand, and T. A. Wilson. J. Physiol. (Lond.) 495:239-246, 1996; A. Legrand, T. A. Wilson, and A. DeTroyer. J. Appl. Physiol. 80:2097-2101, 1996). In the present study, we have tested theprediction further by measuring the response to passive inflation andthe pressure-generating ability of the sternomastoid and scalenemuscles in eight anesthetized dogs. With 1-liter passive inflation, thesternomastoids and scalenes shortened by 2.03 ± 0.17 and 5.98 ± 0.43%, respectively, of their relaxation length(P < 0.001). During maximalstimulation, the two muscles caused similar falls in Pao. However, thesternomastoids had greater mass such that the change in Pao (Pao)per unit muscle mass was 0.19 ± 0.02 cmH₂O/g for the scalenes and only0.07 ± 0.01 cmH₂O/g forthe sternomastoids (P < 0.001).After extension of the neck, there was a reduction in both the muscleshortening during passive inflation and the fall in Pao duringstimulation. The Pao per unit muscle mass was thus closely relatedto the change in length; the slope of the relationship was 3.1. These observations further support the concept that the fractional changes inlength of the respiratory muscles during passive inflation can be usedto predict their pressure-generating ability.

     

Chemical and postural influence on scalene and diaphragmatic activation in humans

The electromyographic activity of the diaphragm (EMGdi) and scalene muscle (EMGsc) was studied in the supine and upright positions, respectively, during hyperoxic progressive hypercapnic rebreathing (HCVR) in five healthy males. End-expiratory esophageal pressure (EEPes) was quantified on a breath-to-breath basis as a reflection of altered end-expiratory lung volume. There was no significant difference in the slopes of EMGdi, expressed as a percentage of maximum at total lung capacity vs. minute volume of ventilation (VI), between the supine and upright positions [0.79 +/- 0.05 (SE) vs. 0.92 +/- 0.17, respectively]. In contrast, the slope of the regression line relating EMGsc to VI was steeper in the upright than in the supine position (0.69 +/- 0.05 vs. 0.35 +/- 0.04, respectively; P less than 0.005). Positive EEPes at comparable VI at the ends of HCVRs were of greater magnitude upright than supine (3.27 +/- 0.68 vs. 4.35 +/- 0.60 cmH2O, respectively, P less than 0.001). We conclude that altering posture has a greater effect on scalene and expiratory muscle activity than on diaphragmatic activity during hypercapnic stimulation.     

Respiratory effects of stimulation of intercostal muscles and saphenous nerve in kittens

Effects of intercostal muscle stimulation were studied in 2- to 7-day-old kittens under ketamine-acepromazine anesthesia. Animals were vagotomized, paralyzed, and artificially ventilated. Stimuli applied during inspiration (TI) inhibited this phase. Stimulus strength necessary for TI inhibition decreased with time. However, an all-or-nothing effect was not always observed. Stimulation during expiration (TE) prolonged this phase. The responsiveness increased with increasing stimulus delay. The effects of intercostal muscle stimulation were compared with those recorded during saphenous nerve stimulation. Stimulation during TI prolonged this phase. Phrenic activity increased after a short-lasting decrease in the on-going activity. Stimulation during the first 50% of TE had variable effects, whereas stimulation with longer delay shortened this phase. Our results indicated that the pattern of breathing in newborns can be affected by both intercostal muscle and other somatic efferents. However, the mechanisms controlling respiratory timing may differ in newborns and in adults. Different effects of respiratory muscle and saphenous nerve stimulation suggest different transmitters involved or different sites of interaction of these inputs with the medullary respiratory rhythm generator.     

Long-term activity in upper- and lower-limb muscles of humans.

Despite limited data on humans, previous studies suggest that there is an association between the duration of daily muscle activity and the proportion of type I muscle fibers. We quantified the activity of limb muscles in healthy men and women during normal use and compared these measurements with published reports on fiber-type proportions. Seven men (age range = 21-28 yr) and seven women (age range = 18-26 yr) participated in two 10-h recording sessions. Electromyogram (EMG) activity of four muscles in nondominant upper (first dorsal interosseus and biceps brachii) and lower limbs (vastus medialis and vastus lateralis) was recorded with surface electrodes. Hand and arm muscles were active for 18% of the recording time, whereas leg muscles were active for only 10% of the recording time. On average, upper-limb muscles were activated 67% more often than lower-limb muscles. When lower-limb muscles were activated, however, the mean amplitude of each burst was greater in leg muscles [18 and 17% maximum voluntary contraction (MVC)] compared with hand (8% MVC) and arm (6% MVC) muscles. Temporal association in activity between pairs of muscles was high for the two lower-limb muscles (r2 = 0.7) and relatively weak for the two upper-limb muscles (r2 = 0.09). Long-term muscle activity was only different between men and women for the biceps brachii muscle. We found no relation between duration of muscle activity in 10-h recordings and the reported values of type I fibers in men and women.     

Respiratory and circulatory activities in carotid body-resected humans.

The respiratory and circulatory activities of patients who underwent carotid body resection (CBR) more than two decades ago were reviewed. No significant ventilatory response to continuous hypoxia was observed. However, in response to stimulation of peripheral chemoreceptors, transient hyperventilation occurred before hypoxemic blood arrived at the central nervous system (single-breath test), which indicated the presence of weak peripheral chemosensitivity. Because of this slight residual peripheral chemosensitivity, which was found shortly after the operation and apparently remained more or less unchanged for greater than 20 years, peripheral chemoreceptor activity, which has been reported in other animal species, does not seem to have returned. Delayed hypoxic hyperventilation reported in dogs and cats with CBR was not observed. Hypoxia significantly depressed the ventilatory response to CO2, but the delayed ventilatory depression with time that has been demonstrated in normal subjects did not occur. In our circulatory studies, hypoxia augmented the heart rate and slightly depressed the stroke volume and total peripheral resistance in the systemic circulation but induced no appreciable changes in arterial blood pressure or cardiac output. We used these results to partition the relative contributions to the overall circulatory response of carotid body stimulation, pulmonary inflation, and other modifying influences. From these calculations, it was inferred that the carotid body reflex plays a dominant role in vascular activities whereas the pulmonary inflation reflex dominates in cardiac activities in humans.     

Aging effects on posture-related modulation of stretch reflex excitability in the ankle muscles in humans

The purpose of this study was to examine the effects of aging on posture-related changes of the stretch reflex excitability in the ankle extensor, soleus (SOL), and flexor, tibialis anterior (TA) muscles. Fourteen neurologically normal elderly (mean 68 ± 6 years) and 12 young (mean 27 ± 3 years) subjects participated. Under two postural conditions, upright standing (STD) and sitting (SIT), stretch reflex electromyographic (EMG) responses in the SOL/TA muscle were elicited by imposing rapid ankle dorsi-/plantar-flexion. Under the SIT condition, subjects were asked to keep the SOL background EMG level, which is identical to that under the STD condition. In the SOL muscle, both groups showed significant enhancement of the short-latency stretch reflex (SLR) response when the posture changed from SIT to STD. In the TA muscle, the young group showed significant enhancement of the middle- (MLR) and long-latency stretch reflex (LLR) when the posture changed from SIT to STD; no such modulation was observed in the elderly group. Since the TA stretch reflex responses under the STD condition were comparable in the young and elderly groups, the lack of posture-related modulation of the TA muscle in the elderly group might be explained by augmented stretch reflex excitability under the SIT condition. The present results suggest that the (1) SOL SLR responses are modulated both in the young and elderly subjects when the posture is changed from SIT to STD, (2) TA MLR and LLR responses are not modulated in the elderly subjects when the posture is changed from SIT to STD, while each response is same between the young and elderly in STD, and (3) the effect of aging on the posture-related stretch reflex differs in the SOL and TA muscles.     

Respiratory function of pharyngeal muscles

This article reviews experimental studies of pharyngeal muscles with emphasis on m. genioglossus as a major muscle dilating pharynx and discusses neuromuscular mechanisms that maintains patency of upper airway. Mechanisms of inspiratory activation of genioglossus muscle in comparative with diaphragm are also discussed. Experimental data suggesting that upper airway muscles have a significant role in compensation of added inspiratory load are presented. It allows to regard pharyngeal dilating muscles as accessory muscles of respiration. Activation of m. genioglossus (together with others muscles dilating the pharynx) decreases airway resistance and thereby facilitates the load compensation function of "pumping" muscles. Similar to diaphragm involvement of the pharynx dilating muscles in the load compensatory response is resulted from a complex integration of several influences originating from mechano- and chemoreceptors.     

Respiratory effects in humans of a 5-day elevation of end-tidal PCO2 by 8 Torr.

The aims of this study were to determine 1) whether ventilatory adaptation occurred over a 5-day exposure to a constant elevation in end-tidal PCO2 and 2) whether such an exposure altered the sensitivity of the chemoreflexes to acute hypoxia and hypercapnia. Ten healthy human subjects were studied over a period of 13 days. Their ventilation, chemoreflex sensitivities, and acid-base status were measured daily before, during, and after 5 days of elevated end-tidal PCO2 at 8 Torr above normal. There was no major adaptation of ventilation during the 5 days of hypercapnic exposure. There was an increase in ventilatory chemosensitivity to acute hypoxia (from 1.35 +/- 0.08 to 1.70 +/- 0.07 l/min/%; P < 0.01) but no change in ventilatory chemosensitivity to acute hypercapnia. There was a degree of compensatory metabolic alkalosis. The results do not support the hypothesis that the ventilatory adaptation to chronic hypercapnia would be much greater with constant elevation of alveolar PCO2 than with constant elevation of inspired PCO2, as has been used in previous studies and in which the feedback loop between ventilation and alveolar PCO2 is left intact.     

Antimutagenic effects in humans.

The application of antimutagenicity studies to human somatic mutation is discussed, with emphasis on the potential for future studies. Five assay-gene combinations are now available for measuring human somatic mutation in lymphocytes and erythrocytes. Results with these combinations have defined the human background levels, and show clear responses of mutant frequency to a variety of mutagens. The testing of antimutagenic effects on background frequencies is feasible, but has not yet been done. The major uncertainty in such studies is the unknown age of mutant cells in the background, since only the newly forming mutants are potentially susceptible to most antimutagenic treatments. Intervention studies in the face of active mutagenicity and the use of other genotoxicity endpoints, such as chromosome aberrations, micronuclei and DNA adducts, are considered briefly.     

Acute effects of thixotropy conditioning of inspiratory muscles on end-expiratory chest wall and lung volumes in normal humans.

Thixotropy conditioning of inspiratory muscles consisting of maximal inspiratory effort performed at an inflated lung volume is followed by an increase in end-expiratory position of the rib cage in normal human subjects. When performed at a deflated lung volume, conditioning is followed by a reduction in end-expiratory position. The present study was performed to determine whether changes in end-expiratory chest wall and lung volumes occur after thixotropy conditioning. We first examined the acute effects of conditioning on chest wall volume during subsequent five-breath cycles using respiratory inductive plethysmography (n = 8). End-expiratory chest wall volume increased after conditioning at an inflated lung volume (P < 0.05), which was attained mainly by rib cage movements. Conditioning at a deflated lung volume was followed by reductions in end-expiratory chest wall volume, which was explained by rib cage and abdominal volume changes (P < 0.05). End-expiratory esophageal pressure decreased and increased after conditioning at inflated and deflated lung volumes, respectively (n = 3). These changes in end-expiratory volumes and esophageal pressure were greatest for the first breath after conditioning. We also found that an increase in spirometrically determined inspiratory capacity (n = 13) was maintained for 3 min after conditioning at a deflated lung volume, and a decrease for 1 min after conditioning at an inflated lung volume. Helium-dilution end-expiratory lung volume increased and decreased after conditioning at inflated and deflated lung volumes, respectively (both P < 0.05; n = 11). These results suggest that thixotropy conditioning changes end-expiratory volume of the chest wall and lung in normal human subjects.     

Respiratory responses induced by the activation of somatic nociceptive afferents in humans.

To further investigate the role of somatic nociceptive afferents in the neural control of breathing, we studied the respiratory effects of their activation by means of either electrical stimulation or ischemic pain in 14 healthy volunteers. Painful electrical cutaneous stimulation increased respiratory frequency (f), mean inspiratory flow (VT/TI), and rate of rise (XP/TI) of integrated electromyographic activity of diaphragm (IEMGdi). Painful muscular electrical stimulation caused similar but larger changes accompanied by increases in tidal volume (VT), peak XP of IEMGdi, and ventilation (VE); it also entrained respiratory rhythm. Ischemic pain, which was characterized by a progressively increasing intensity, caused augmentation in respiratory activity that displayed an increasing trend: VE, f, VT, XP, VT/TI, and XP/TI increased. In the light of available literature, it seems conceivable to suggest that respiratory responses to painful electrical stimulation are mediated through the activation of cutaneous (A delta) and muscular (group III) fine-myelinated afferents, and responses to ischemic pain are mediated by the activation of both fine myelinated (group III) and unmyelinated (group IV) muscular afferents. The input conveyed by these afferents may constitute an effective stimulus to respiration in humans.     

Activity of respiratory muscles in upright and recumbent humans

It is established that during tidal breathing the rib cage expands more than the abdomen in the upright posture, whereas the reverse is usually true in the supine posture. To explore the reasons for this, we studied nine normal subjects in the supine, standing, and sitting postures, measuring thoracoabdominal movement with magnetometers and respiratory muscle activity via integrated electromyograms. In eight of the subjects, gastric and esophageal pressures and diaphragmatic electromyograms via esophageal electrodes were also measured. In the upright postures, there was generally more phasic and tonic activity in the scalene, sternocleidomastoid, and parasternal intercostal muscles. The diaphragm showed more phasic (but not more tonic) activity in the upright postures, and the abdominal oblique muscle showed more tonic (but not phasic) activity in the standing posture. Relative to the esophageal pressure change with inspiration, the inspiratory gastric pressure change was greater in the upright than in the supine posture. We conclude that the increased rib cage motion characteristic of the upright posture owes to a combination of increased activation of rib cage inspiratory muscles plus greater activation of the diaphragm that, together with a stiffened abdomen, acts to move the rib cage more effectively.     

Respiratory muscle injury in animal models and humans

Respiratory muscle injury may result from excessive loading due to a decrease in respiratory muscle strength, an increase in the work of breathing, or an increase in the rate of ventilation. Other conditions such as hypoxemia, hypercapnia, aging, decreased nutrition, and immobilization may potentiate respiratory muscle injury. Respiratory muscle injury has been shown in animal models using direct muscle or phrenic nerve stimulation, acute inspiratory resistive loading, tracheal banding, corticosteroids, phrenic nerve section, and the mdx mouse. Although numerous examples of diaphragm injury have been shown in animal models, evidence in humans is sparse. Potential mechanisms which may contribute to respiratory muscle injury include high levels of intracellular calcium-activated degradative enzymes, non-uniformity of stresses and strains, plasma membrane disruptions, and activation of the inflammatory process.     

Influence of long-term spaceflight on neuromechanical properties of muscles in humans.

Reflex and elastic properties of the triceps surae (TS) were measured on 12 male cosmonauts 28-40 days before a 3- to 6-mo spaceflight, 2 or 3 days after return (R+2/+3) and a few days later (R+5/+6). H reflexes to electrical stimulations and T reflexes to tendon taps gave the reflex excitability at rest. Under voluntary contractions, reflex excitability was assessed by the stretch reflex, elicited by sinusoidal length perturbations. Stiffness measurements concerned the musculoarticular system in passive conditions and the musculotendinous complex in active conditions. Results indicated 1) no changes (P > 0.05) in H reflexes, whatever the day of test, and 2) increase in T reflexes (P < 0.05) by 57%, despite a decrease (P < 0.05) in musculoarticular stiffness (11%) on R+2/+3. T reflexes decreased (P < 0.05) between R+2/+3 and R+5/+6 (-21%); 3) increase in stretch reflexes (P < 0.05) on R+2/+3 by 31%, whereas it decreased (P < 0.05) between R+2/+3 and R+5/+6 (-29%). Musculotendinous stiffness was increased (P < 0.05) whatever the day of test (25%). Links between changes in reflex and stiffness were also studied by considering individual data. At R+2/+3, correlated changes between T reflexes and musculoarticular stiffness suggested that, besides central adaptive phenomena, musculoarticular structures took part in the reflex adaptation. This mechanical contribution was confirmed when data collected at R+2/+3 and R+5/+6 were used because correlations between changes in stretch reflexes and musculotendinous stiffness were improved. In conclusion, the present study shows that peripheral influences take part in reflex changes in gravitational unloaded muscles, but can only be revealed when central influences are reduced.