Abdominal muscle use during breathing in unanesthetized dogs

The pattern of abdominal muscle use during breathing in unanesthetized dogs is unknown. Therefore, we have recorded the electromyograms of the rectus abdominis, external oblique, and transversus abdominis in eight conscious animals breathing quietly in the sitting, standing, and prone postures. During quiet breathing in the sitting posture, all animals invariably had a large amount of phasic expiratory activity in the transversus abdominis. In contrast, only four animals showed some expiratory activity in the external oblique, and only one animal had expiratory activity in the rectus abdominis. A similar pattern was observed when the animals were standing or lying prone, although the amount of expiratory activity was less in this posture. Bilateral cervical vagotomy in four animals did not affect the degree of transversus abdominis expiratory activation or the influence of posture. We conclude that in conscious dogs 1) the abdominal muscles play an important role during breathing and make spontaneous quiet expiration a very active process, 2) the transversus abdominis is the primary respiratory muscle of the abdomen, and 3) unlike in anesthetized animals, extrapulmonary receptors play a major role in promoting abdominal expiratory contraction.     

Breathing strategy of the adult horse (Equus caballus) at rest

To investigate the mechanism underlying the polyphasic airflow pattern of the equine species, we recorded airflow, tidal volum, rib cage and abdominal motion, and the sequence of activation of the diaphragm, intercostal, and abdominal muscles during quiet breathing in nine adult horses standing at rest. In addition, esophageal, abdominal, and transdiaphragmatic pressures were simultaneously recorded using balloon-tipped catheters. Analysis of tidal flow-volume loops showed that, unlike humans, the horse at rest breathes around, rather than from, the relaxed volume of the respiratory system (Vrx). Analysis of the pattern of electromyographic activities and changes in generated pressures during the breathing cycle indicate that the first part of expiration is passive, as in humans, with deflation toward Vrx, but subsequent abdominal activity is responsible for a second phase of expiration: active deflation to below Vrx. From this end-expiratory volume, passive inflation occurs toward Vrx, followed by a second phase of inspiration: active inflation to above Vrx, brought about by inspiratory muscle contraction. Under these conditions the abdominal muscles appear to share the principal pumping duties with the diaphragm. Adoption of this breathing strategy by the horse may relate to its peculiar thoracoabdominal anatomic arrangement and to its very low passive chest wall compliance. We conclude that there is a passive and active phase to both inspiration and expiration due to the coordinated action of the respiratory pump muscles responsible for the resting adult horse's biphasic inspiratory and expiratory airflow pattern. This unique breathing pattern perhaps represents a strategy of minimizing the high elastic work of breathing in this species, at least at resting breathing frequencies.     

Chest wall motion during epidural anesthesia in dogs

To determine the relative contribution of rib cage and abdominal muscles to expiratory muscle activity during quiet breathing, we used lumbar epidural anesthesia in six pentobarbital sodium-anesthetized dogs lying supine to paralyze the abdominal muscles while leaving rib cage muscle motor function substantially intact. A high-speed X-ray scanner (Dynamic Spatial Reconstructor) provided three-dimensional images of the thorax. The contribution of expiratory muscle activity to tidal breathing was assessed by a comparison of chest wall configuration during relaxed apnea with that at end expiration. We found that expiratory muscle activity was responsible for approximately half of the changes in thoracic volume during inspiration. Paralysis of the abdominal muscles had little effect on the pattern of breathing, including the contribution of expiratory muscle activity to tidal breathing, in most dogs. We conclude that, although there is consistent phasic expiratory electrical activity in both the rib cage and the abdominal muscles of pentobarbital-anesthetized dogs lying supine, the muscles of the rib cage are mechanically the most important expiratory muscles during quiet breathing.     

The effect of posture on respiratory activity of the abdominal muscles

The purpose of this study was to determine the influence of posture on the expiratory activity of the abdominal muscles. Fifteen young adult men participated in the study. Activities of the external oblique abdominis, internal oblique abdominis, and rectus abdominis muscles were measured electromyographically in various postures. We used a pressure threshold in order to activate the abdominal muscles as these muscles are silent at rest. A spirometer was used to measure the lung volume in various postures. Subjects were placed in the supine, standing, sitting, and sitting-with-elbow-on-the-knee (SEK) positions. Electromyographic activity and mouth pressure were measured during spontaneous breathing and maximal voluntary ventilation under the respiratory load. We observed that the lung volume changed with posture; however, the breathing pattern under respiratory load did not change. During maximal voluntary ventilation, internal oblique abdominis muscle expiratory activity was lower in the SEK position than in any other position, external oblique abdominis muscle inspiratory activity was lower in the supine position than in any other position, and internal oblique abdominis muscle activity was higher in the standing position than in any other position. During spontaneous breathing, external oblique abdominis muscle activity was higher during expiration and inspiration in the SEK position than in any other position. The internal oblique abdominis muscle activity was higher during both inspiration and expiration in the standing position than in any other position. The rectus abdominis muscle activity did not change with changes in posture during both inspiration and expiration. Increase in the external oblique abdominis activity in the SEK position was due to anatomical muscle arrangement that was consistent with the direction of lower rib movement. On the other hand, increase in the internal oblique abdominis activity in the standing position was due to stretching of the abdominal wall by the viscera. We concluded that differences in activity were due to differences in the anatomy of the abdominal muscles and the influence of gravity.     

Relative contributions of rib cage and abdomen to breathing in normal subjects.

By use of the method of Konno and Mead and the respiratory magnetometer, the partition of respired gas volumes into rib cage and diaphragm-abdomen components was accomplished in 81 normal subjects including 32 young and middle-aged men, 29 young and middle-aged women, and 20 elderly men. Studied were isovolume maneuvers and the relaxation configuration over the inspiratory capacity range, quiet tidal breathing, increased amplitudes of slow breathing, rapid inspirations and expirations, and both quiet and forceful phonation. No major differences were noted between men and women or between the young and the elderly during any respiratory acts. During quiet breathing most normal subjects are abdominal breathers when supine and thoracic breathers when upright. Rapid respiratory maneuvers were accomplished mostly through rib cage displacement suggesting that rib cage muscles are capable of more rapid action than diaphragm and abdominal muscles. Data from deep breathing and rapid maneuvers supported the view that abdominal and rib cage muscles often act to optimize the mechanical (length-tension) advantage of the diaphragm.     

Changes in cardiac output during sustained maximal ventilation in humans

To determine the increment in cardiac output and in O2 consumption (Vo2) from quiet breathing to maximal sustained ventilation, Vo2 and cardiac output were measured using an acetylene rebreathing technique in five subjects. Cardiac output and Vo2 were measured multiple times in each subject at rest and during sustained maximal ventilation. During maximal ventilation subjects breathed 5% CO2 to prevent hypocapnia. The increase in cardiac output from rest to maximal breathing was taken as an estimate of respiratory muscle blood flow and was used to calculate the arteriovenous O2 content difference across the respiratory muscles from the Fick equation. Cardiac output increased by 4.3 +/- 1.0 l/min (mean +/- SD), from 5.6 +/- 0.7 l/min at rest to 9.9 +/- 1.1 l/min, during maximal ventilations ranging from 127 to 193 l/min. Vo2 increased from 312 +/- 29 to 723 +/- 69 ml/min during maximal ventilation. O2 extraction across the respiratory muscles during maximal breathing was 9.6 +/- 1.0 vol% (range 8.5 to 10.7 vol%). These values suggest an upper limit of respiratory muscle blood flow of 3-5 l/min during unloaded maximal sustained ventilation.     

Effects of midline laparotomy on expiratory muscle activation in anesthetized dogs

Abdominal surgery has a marked inhibitory influence on the diaphragm, but its effect on the expiratory muscles is not known. Therefore, we have recorded the electromyograms of the triangularis sterni, abdominal external oblique, and transversus abdominis before and after a midline laparotomy in 10 anesthetized, spontaneously breathing dogs. Measurements were obtained during quiet breathing in the supine posture, during breathing against expiratory threshold loads, during head-up tilting, and during hyperoxic hypercapnia. Expiratory activation of the transversus abdominis in all conditions was considerably reduced after laparotomy. This reduction was real, as no change in the compound muscle action potential during single pulse stimulation was observed. In contrast, expiratory recruitment of either the triangularis sterni or the abdominal external oblique was maintained or increased. We therefore conclude that laparotomy inhibits not only activation of the diaphragm during inspiration but also activation of the transversus abdominis during expiration. Visceral afferents thus affect in concert the two respiratory muscles lining the peritoneum. The present findings also emphasize the important fact that the pattern of activation of a particular abdominal muscle is not necessarily representative of the entire abdominal musculature.     

A Closed-Loop Model of the Respiratory System: Focus on Hypercapnia and Active Expiration

Breathing is a vital process providing the exchange of gases between the lungs and atmosphere. During quiet breathing, pumping air from the lungs is mostly performed by contraction of the diaphragm during inspiration, and muscle contraction during expiration does not play a significant role in ventilation. In contrast, during intense exercise or severe hypercapnia forced or active expiration occurs in which the abdominal “expiratory” muscles become actively involved in breathing. The mechanisms of this transition remain unknown. To study these mechanisms, we developed a computational model of the closed-loop respiratory system that describes the brainstem respiratory network controlling the pulmonary subsystem representing lung biomechanics and gas (O₂ and CO₂) exchange and transport. The lung subsystem provides two types of feedback to the neural subsystem: a mechanical one from pulmonary stretch receptors and a chemical one from central chemoreceptors. The neural component of the model simulates the respiratory network that includes several interacting respiratory neuron types within the Bötzinger and pre-Bötzinger complexes, as well as the retrotrapezoid nucleus/parafacial respiratory group (RTN/pFRG) representing the central chemoreception module targeted by chemical feedback. The RTN/pFRG compartment contains an independent neural generator that is activated at an increased CO₂ level and controls the abdominal motor output. The lung volume is controlled by two pumps, a major one driven by the diaphragm and an additional one activated by abdominal muscles and involved in active expiration. The model represents the first attempt to model the transition from quiet breathing to breathing with active expiration. The model suggests that the closed-loop respiratory control system switches to active expiration via a quantal acceleration of expiratory activity, when increases in breathing rate and phrenic amplitude no longer provide sufficient ventilation. The model can be used for simulation of closed-loop control of breathing under different conditions including respiratory disorders.     

Electromyographic activity of expiratory muscles in the rat

We examined the participation of expiratory muscles on breathing in the rat. The experiments were performed on 16 male rats in halothane [1.5%] or urethane [1.3 g/kg i.p.] anaesthesia. We recorded the electromyographic [EMG] activity of intercostal and abdominal muscles with a concentric needle electrode during quiet breathing, breathing against increased pressure in the airways and during the expiration reflex. In halothane anaesthesia the EMG expiratory phasic activity was observed only in internal intercostal muscles in 40% of spots examined during quiet breathing and in 58.5% when breathing against increased pressure. The EMG activity during the expiratory reflex was difficult to evaluate. In the abdominal muscles permanent EMG activity was found in 66% of trials. In urethane anaesthesia no phasic expiratory EMG activity was observed in intercostal or abdominal muscles. In abdominal muscles in 9% of trials a permanent activity was found.     

Superextension and supercontraction in locust ovipositor muscles

A ten times elongation of certain abdominal intersegmental muscles occurs in female locusts during digging prior to oviposition. During and after oviposition the muscles contract, shortening by up to 90% or more, restoring the resting positions of the abdominal segments.Discontinuous Z-discs permit supercontraction at the resting length and then fragment into Z-bodies when the muscle is stretched, so enabling it to superextend without loss of the contractile property. In this superextended state the fibres resemble smooth muscles. After oviposition, the muscle fibres contract but the sarcomeres are not restored completely, some of the Z-bodies being unevenly distributed in the recontracted fibres. Locust ovipositor muscle has the most extreme example of Z-disc disagregation known from the insects and is the insect muscle which approaches most closely the smooth muscle condition.Two types of motor nerve innervate this muscle, one is ordinary and the other, containing granules, resembles an octopaminergic fibre possibly involved in regulating a catch mechanism in the muscle.The physiological requirements for egg-laying with an extensible ovipositor, which is also part of the normally functioning abdomen, are well met by the ultrastructural specializations of locust ovipositor muscles.     

Increased ventilation does not impair maximal voluntary contractions of the elbow flexors.

Exercise performance is impaired by increased respiratory work, yet the mechanism for this is unclear. This experiment assessed whether neural drive to an exercising muscle was affected by cortically driven increases in ventilation. On each of 5 days, eight subjects completed a 2-min maximal voluntary contraction (MVC) of the elbow flexor muscles, followed by 4 min of recovery, while transcranial magnetic stimulation tested for suboptimal neural drive to the muscle. On 1 day, subjects breathed without instructions under normocapnia. During the 2-min MVC, ventilation was approximately 3.5 times that at rest. On another day, subjects breathed without instruction under hypercapnia. During the 2-min MVC, ventilation was approximately 1.5 times that on the normocapnic day. On another 2 days under normocapnia, subjects voluntarily matched their breathing to the uninstructed breathing under normocapnia and hypercapnia using target feedback of the rate and inspiratory volume. On a fifth day under normocapnia, the volume feedback was set to each subject's vital capacity. On this day, ventilation during the 2-min MVC was approximately twice that on the uninstructed normocapnic day (or approximately 7 times rest). The experimental manipulations succeeded in producing voluntary and involuntary hyperpnea. However, maximal voluntary force, fatigue and voluntary activation of the elbow flexor muscles were unaffected by cortically or chemically driven increases in ventilation. Results suggest that any effects of increased respiratory work on limb exercise performance are not due to a failure to drive both muscle groups optimally.     

Voluntary hyperventilation changes recruitment order of parasternal intercostal motor units

The order of recruitment of single-motor units in parasternal intercostal muscles during inspiration was studied in normal human subjects during quiet breathing and voluntary hyperventilation. Electromyograms were recorded from the second and third intercostal spaces by means of bipolar fine wire electrodes. Flow at the mouth, volume, end-expired CO2, and rib cage and abdominal anterior-posterior diameters were monitored. Single-motor units were identified using criteria of amplitude and shape, and the time of first appearance of each unit in each inspiration was noted. Hyperventilation was performed with visual feedback of the display of rib cage and abdomen excursions, keeping the ratio of rib cage to abdominal expansion. Subjects were normocapnic in quiet breathing and developed hypocapnia during hyperventilation. Recruitment order was stable in quiet breathing, but in some cases was altered during voluntary hyperventilation. Some low threshold units that fired early in the breath in quiet breathing fired earlier at the beginning of a period of voluntary hyperventilation but progressively later in the breath as hyperventilation went on, whereas later firing units moved progressively toward the early part of inspiration. This suggests that different groups of motoneurons in the pool supplying parasternal intercostal muscles receive different patterns of synaptic input.     

Triangularis sterni muscle use in supine humans

The electrical activity of the triangularis sterni (transversus thoracis) muscle was studied in supine humans during resting breathing and a variety of respiratory and nonrespiratory maneuvers known to bring the abdominal muscles into action. Twelve normal subjects, of whom seven were uninformed and untrained, were investigated. The electromyogram of the triangularis sterni was recorded using a concentric needle electrode, and it was compared with the electromyograms of the abdominal (external oblique and rectus abdominis) muscles. The triangularis sterni was usually silent during resting breathing. In contrast, the muscle was invariably activated during expiration from functional residual capacity, expulsive maneuvers, "belly-in" isovolume maneuvers, static head flexion and trunk rotation, and spontaneous events such as speech, coughing, and laughter. When three trained subjects expired voluntarily with considerable recruitment of the triangularis sterni and no abdominal muscle activity, rib cage volume decreased and abdominal volume increased. These results indicate that unlike in the dog, spontaneous quiet expiration in supine humans is essentially a passive process; the human triangularis sterni, however, is a primary muscle of expiration; and its neural activation is largely coupled with that of the abdominals. The triangularis sterni probably contributes to the deflation of the rib cage during active expiration.     

Activity in torso muscles during relaxed standing

Electromyographic activity of erector spinae, external oblique, and rectus abdominis muscles was studied during relaxed standing compared to lying down. Activity in the forearm extensors and forearm flexors was also studied. Surface electrodes were used. Each of the torso muscles exhibited 0.2 microV of activity and the forearm muscles 0.1 microV while subjects were relaxed and lying down. During quiet standing the erector spinae, external oblique, and rectus abdominis muscles showed a median activity of 1.0 microV, 2.5 microV, and 0.7 microV respectively (for a minimum of ten 10-sec samples per subject). Examination of the integrated records during standing revealed no periods without increased muscle activity in the torso muscles. By contrast, activity in the forearm muscles did not increase during standing. The major superficial muscles of posture in the torso appear to act as guy wires, being continually active during standing. There is no support for hypotheses of passive support for the torso, nor do torso muscles act in either/or fashion; both anterior and posterior muscles are active at once. There is no sign of generally increased muscle tone in all muscles or in extensors; only the postural muscles are continuously active.     

Reproducibility and responsiveness of a noninvasive EMG technique of the respiratory muscles in COPD patients and in healthy subjects.

In the present study, we assessed the reproducibility and responsiveness of transcutaneous electromyography (EMG) of the respiratory muscles in patients with chronic obstructive pulmonary disease (COPD) and healthy subjects during breathing against an inspiratory load. In seven healthy subjects and seven COPD patients, EMG signals of the frontal and dorsal diaphragm, intercostal muscles, abdominal muscles, and scalene muscles were derived on 2 different days, both during breathing at rest and during breathing through an inspiratory threshold device of 7, 14, and 21 cm H2O. For analysis, we used the logarithm of the ratio of the inspiratory activity during the subsequent loads and the activity at baseline [log EMG activity ratio (EMGAR)]. Reproducibility of the EMG was assessed by comparing the log EMGAR values measured at test days 1 and 2 in both groups. Responsiveness (sensitivity to change) of the EMG was assessed by comparing the log EMGAR values of the COPD patients to those of the healthy subjects at each load. During days 1 and 2, log EMGAR values of the diaphragm and the intercostal muscles correlated significantly. For the scalene muscles, significant correlations were found for the COPD patients. Although inspiratory muscle activity increased significantly during the subsequent loads in all participants, the COPD patients displayed a significantly greater increase in intercostal and left scalene muscle activity compared with the healthy subjects. In conclusion, the present study showed that the EMG technique is a reproducible and sensitive technique to assess breathing patterns in COPD patients and healthy subjects.     

Diaphragm length adjustments with body position changes in the awake dog

Sonomicrometry was used to measure end-expiratory length and tidal shortening of the costal and crural diaphragm in awake chronically instrumented dogs in the right lateral decubitus, standing, and sitting postures. End-expiratory length did not change significantly in standing but fell by 11.5% for the costal and by 14.4% for the crural segment in sitting, when compared with decubitus position. Tidal shortening of both segments did not change significantly in the three postures. From decubitus to sitting, diaphragmatic electromyogram (EMG) activity increased only in some dogs, not significantly for the group. The inspiratory swing of abdominal pressure was always positive in decubitus and negative in standing and sitting. In the latter two postures, abdominal pressure increased gradually during expiration and fell in inspiration, suggesting a phasic expiratory contraction of abdominal muscles. We conclude that diaphragmatic tidal shortening is maintained in the different postures assumed by the awake dog during resting breathing. It seems that the main compensatory mechanism for changes in diaphragmatic operational length is a phasic expiratory contraction of the abdominal muscles rather than an increase in diaphragmatic EMG activity.     

Impedance of the chest wall during sustained respiratory muscle contraction

We measured total chest wall impedance (Zw), "pathway impedances" of the rib cage (Zrcpath), and diaphragm-abdomen (Zd-apath), and impedance of the belly wall including abdominal contents (Zbw+) in five subjects during sustained expiratory (change in average pleural pressure [Ppl] from relaxation = 10 and 20 cmH2O) and inspiratory (change in Ppl = -10 and -20 cmH2O) muscle contraction, using forced oscillatory techniques (0.5-4 Hz) we have previously reported for relaxation (J. Appl. Physiol. 66: 350-359, 1989). Chest wall configuration and mean lung volume were kept constant. Zw, Zrcpath, Zd-apath, and Zbw+ all increased greatly at each frequency during expiratory muscle contraction; increases were proportional to effort. Zw, Zrcpath, and Zd-apath increased greatly during inspiratory muscle contraction, but Zbw+ did not. Resistances and elastances calculated from each of the impedances showed the same changes during muscle contraction as the corresponding impedances. Each of the resistances decreased as frequency increased, independent of effort; elastances generally increased with frequency. These frequency dependencies were similar to those measured in relaxed or tetanized isolated muscle during sinusoidal stretching (P.M. Rack, J. Physiol. Lond. 183: 1-14, 1966). We conclude that during respiratory muscle contraction 1) chest wall impedance increases, 2) changes in regional chest wall impedances can be somewhat independent, depending on which muscles contract, and 3) increases in chest wall impedance are due, at least in part, to changes in the passive properties of the muscles themselves.     

Protons act as a transmitter for muscle contraction in C. elegans

Muscle contraction is normally mediated by the release of neurotransmitters from motor neurons. Here we demonstrate that protons can act as a direct transmitter from intestinal cells to stimulate muscle contraction. During the C. elegans defecation motor program the posterior body muscles contract even in the absence of neuronal inputs or vesicular neurotransmission. In this study, we demonstrate that the space between the intestine and the muscle is acidified just prior to muscle contraction and that the release of caged protons is sufficient to induce muscle contraction. PBO-4 is a putative Na+/H+ ion exchanger expressed on the basolateral membrane of the intestine, juxtaposed to the posterior body muscles. In pbo-4 mutants the extracellular space is not acidified and the muscles fail to contract. The pbo-5 and pbo-6 genes encode subunits of a "cys-loop" proton-gated cation channel required for muscles to respond to acidification. In heterologous expression assays the PBO receptor is half-maximally activated at a pH of 6.8. The identification of the mechanisms for release and reception of proton signals establishes a highly unusual mechanism for intercellular communication.     

Posture effects on timing of abdominal muscle activity during stimulated ventilation.

In humans during stimulated ventilation, substantial abdominal muscle activity extends into the following inspiration as postexpiratory expiratory activity (PEEA) and commences again during late inspiration as preexpiratory expiratory activity (PREA). We hypothesized that the timing of PEEA and PREA would be changed systematically by posture. Fine-wire electrodes were inserted into the rectus abdominis, external oblique, internal oblique, and transversus abdominis in nine awake subjects. Airflow, end-tidal CO2, and moving average electromyogram (EMG) signals were recorded during resting and CO2-stimulated ventilation in both supine and standing postures. Phasic expiratory EMG activity (tidal EMG) of the four abdominal muscles at any level of CO2 stimulation was greater while standing. Abdominal muscle activities during inspiration, PEEA, and PREA, were observed with CO2 stimulation, both supine and standing. Change in posture had a significant effect on intrabreath timing of expiratory muscle activation at any level of CO2 stimulation. The transversus abdominis showed a significant increase in PEEA and a significant decrease in PREA while subjects were standing; similar changes were seen in the internal oblique. We conclude that changes in posture are associated with significant changes in phasic expiratory activity of the four abdominal muscles, with systematic changes in the timing of abdominal muscle activity during early and late inspiration.     

Activation of the parasternal intercostals during breathing efforts in human subjects

We have tested the possibility that the electromyographic (EMG) activity present in the parasternal intercostal muscles during quiet inspiration was reflexive, rather than agonistic, in nature. Using concentric needle electrodes we measured parasternal EMG activity in four normal subjects during various inspiratory maneuvers. We found that 1) phasic inspiratory activity was invariably present in the parasternal intercostals during quiet breathing, 2) the parasternal EMG activity was generally increased during attempts to perform the tidal breathing maneuver with the diaphragm alone, 3) parasternal EMG activity was markedly decreased or suppressed in the presence of rib cage distortion during diaphragmatic isovolume maneuvers, and 4) that EMG activity could not be voluntarily suppressed during breathing unless the inspired volume was trivial. We conclude that the parasternal EMG activity detected during quiet inspiration in the normal subjects depends on a central involuntary mechanism and is not related to activation of intercostal mechanoreceptors.