Effect of chest wall vibration on the canine diaphragm during breathing.

High-frequency mechanical vibration of the ribcage reduces dyspnoea in patients with chronic obstructive pulmonary disease, and the suggestion has been made that this effect might be related to a decrease in central respiratory drive resulting from an increase in afferent inputs from intercostal muscles. In the present studies, the effects of ribcage vibration on central respiratory drive have been assessed without the confounding influence of conscious reactions. The electromyographic (EMG) activity of the diaphragm and the changes in pleural (Ppl) and abdominal (Pab) pressure were measured in six anaesthetized, spontaneously-breathing dogs while the rostral, the middle, or the caudal portion of the ribcage was vibrated at intervals during inspiration. The EMG activity of the external and parasternal intercostals was also measured. Ribcage vibration consistently elicited a marked increase in the inspiratory EMG activity recorded from the external intercostals, thus indicating that the procedure did activate intercostal muscle spindles. However, no alteration in diaphragmatic or parasternal intercostal EMG activity was seen in any animal. Transdiaphragmatic pressure and the relationship between deltaPab and deltaPpl during inspiration were also unaltered. The authors conclude that ribcage vibration and, with it, stimulation of external intercostal muscle spindles has no significant influence on phrenic motoneurones or on medullary inspiratory neurones. It is unlikely, therefore, that the beneficial effect of the procedure on dyspnoea results from a specific reduction in central respiratory drive.

[The pneumology department]. / Le Service de Pneumologie.

The department of pneumology of the Erasme hospital exists since 25 years. The basic clinical activities include pulmonary function testing (7,500 patients per year), endoscopy, including interventional endoscopy (1,500 patients per year), thoracic oncology, allergology, rehabilitation and aid to smoking cessation. The following expertise fields have been largely developed: lung transplantation, treatment of cystic fibrosis in collaboration with the children's hospital Reine Fabiola, occupational.

Contribution of spindle reflexes to post-inspiratory activity in the canine external intercostal muscles.

1. The external intercostal muscles have greater post-inspiratory activity than the parasternal intercostal muscles and are more abundantly supplied with muscle spindles. In the present study, the hypothesis was tested that spindle afferent inputs play a major role in determining this activity. 2. The electrical activity of the external and parasternal intercostal muscles in the rostral interspaces was recorded in anaesthetized spontaneously breathing dogs, and the ribs were manipulated so as to alter their normal caudal displacement and the normal lengthening of the muscles in early expiration. 3. Post-inspiratory activity in the external intercostal muscles showed a reflex decrease when the caudal motion of the ribs and the lengthening of the muscles was impeded, and it showed a reflex increase when the rate of caudal rib motion and muscle lengthening was increased. In contrast, the small post-inspiratory activity in the parasternal intercostal muscles remained unchanged. 4. When the two ribs making up the interspace investigated were locked to keep muscle length constant, post-inspiratory activity in the external intercostal muscles was reduced and no longer responded to cranial rib manipulation. 5. These observations confirm that afferent inputs from muscle receptors, presumably muscle spindles, are a primary determinant of post-inspiratory activity in the canine external intercostal muscles. In anaesthetized animals, the contribution of central control mechanisms to this activity is small.

Response of the canine internal intercostal muscles to chest wall vibration.

Although high-frequency mechanical vibration of the rib cage reduces dyspnea, its effects on the respiratory muscles are largely unknown. We have previously shown that in anesthetized dogs, vibrating the rib cage during inspiration elicits a marked increase in the inspiratory electromyographic (EMG) activity recorded from the external intercostal muscles but does not affect tidal volume (VT). In the present studies, we have tested the hypothesis that the maintenance of VT results from the concomitant contraction of the internal interosseous (expiratory) intercostals. When the rib cage was vibrated (40 Hz) during hyperventilation-induced apnea, a prominent activity was recorded from the external intercostals but no activity was recorded from the internal intercostals, including when the muscles were lengthened by passive inflation. The internal intercostals remained also silent when vibration was applied during spontaneous inspiration, and the phasic expiratory EMG activity recorded from them was unaltered when vibration was applied during expiration. Thus, the internal interosseous intercostals in dogs are much less sensitive to vibration than the external intercostals, and they do not interfere with the action of these latter during rib cage vibration. This lack of sensitivity might be the result of a reflex inhibition of the muscle spindle afferents by afferents from external intercostal muscle spindles.

Respiratory effects of the external and internal intercostal muscles in humans.

The current conventional view of intercostal muscle actions is based on the theory of Hamberger (1749) and maintains that as a result of the orientation of the muscle fibres, the external intercostals have an inspiratory action on the lung and the internal interosseous intercostals have an expiratory action. Recent studies in dogs, however, have shown that this notion is only approximate. In the present studies, the respiratory actions of the human external and internal intercostal muscles were evaluated by applying the Maxwell reciprocity theorem. Thus the orientation of the muscle fibres relative to the ribs and the masses of the muscles were first assessed in cadavers. Five healthy individuals were then placed in a computed tomographic scanner to determine the geometry of the ribs and their precise transformation during passive inflation to total lung capacity. The fractional changes in length of lines with the orientation of the muscle fibres were then computed to obtain the mechanical advantages of the muscles. These values were finally multiplied by muscle mass and maximum active stress (3.0 kg cm-2) to evaluate the potential effects of the muscles on the lung. The external intercostal in the dorsal half of the second interspace was found to have a large inspiratory effect. However, this effect decreases rapidly in the caudal direction, in particular in the ventral portion of the ribcage. As a result, it is reversed into an expiratory effect in the ventral half of the sixth and eighth interspaces. The internal intercostals in the ventral half of the sixth and eighth interspaces have a large expiratory effect, but this effect decreases dorsally and cranially. The total pressure generated by all the external intercostals during a maximum contraction would be -15 cmH2O, and that generated by all the internal interosseous intercostals would be +40 cmH2O. These pressure changes are substantially greater than those induced by the parasternal intercostal and triangularis sterni muscles, respectively.

Heterogeneity of metabolic activity in the canine parasternal intercostals during breathing.

In the dog, the inspiratory mechanical advantage of the parasternal intercostals shows a marked spatial heterogeneity, whereas the expiratory mechanical advantage of the triangularis sterni is relatively uniform. The contribution of a particular respiratory muscle to lung volume expansion during breathing, however, depends both on the mechanical advantage of the muscle and on its neural input. To evaluate the distribution of neural input across the canine parasternal intercostals and triangularis sterni, we have examined the distribution of metabolic activity among these muscles in seven spontaneously breathing animals by measuring the uptake of the glucose tracer analog [(18)F]fluorodeoxyglucose (FDG). FDG uptake in any given parasternal intercostal was greatest in the medial bundles and decreased rapidly toward the costochondral junctions. In addition, FDG uptake in the medial parasternal bundles increased from the first to the second interspace, plateaued in the second through fifth interspaces, and then decreased progressively toward the eighth interspace. In contrast, uptake in the triangularis sterni showed no significant rostrocaudal gradient. These results overall strengthen the idea that the spatial distribution of neural input within a particular set of respiratory muscles is closely matched with the spatial distribution of mechanical advantage.

Response of the rabbit diaphragm to tendon vibration.

To evaluate the potential role of diaphragmatic muscle spindles in the act of breathing, we have recorded the electromyograms of the diaphragm and the external intercostal muscle in the third interspace during high-frequency mechanical vibration (50 Hz) of the central tendon in eight anesthetized, spontaneously breathing rabbits. Vibration induced a consistent, clear-cut increase in the inspiratory activity recorded from the external intercostal, thus indicating that the mechanical stimulus applied to the diaphragm was strong enough to trigger muscle spindles at distant sites. However, vibration did not elicit any alteration in costal or crural diaphragmatic activity in any animal. Similarly, when vibration was applied during hyperventilation-induced apnea, activity was recorded in the external intercostal but not in the diaphragm. These observations support the traditional view that the diaphragm is poorly endowed with muscle spindles and that these play little or no significant role in the act of breathing.

Regional differences in serotonergic input to canine parasternal intercostal motoneurons.

There is a mediolateral gradient in activation of the parasternal intercostal (PI) muscle during inspiration. In the present study, we tested the hypotheses that serotonergic [5-hydroxytryptamine (5-HT)] input from descending central drive and/or intrinsic size-related properties of PI motoneurons leads to the differential activation of PI muscles. In dogs, PI motoneurons innervating the medial and lateral regions of the PI muscles at the T(3)-T(5) interspaces were retrogradely labeled by intramuscular injection of cholera toxin B subunit. After a 10-day survival period, PI motoneurons and 5-HT terminals were visualized by using immunohistochemistry and confocal imaging. There were no differences in motoneuron morphology among motoneurons innervating the medial and lateral regions of the PI muscle. However, the number of 5-HT terminals and the 5-HT terminal density (normalized for surface area) were greater in motoneurons innervating the medial region of the PI muscle compared with the lateral region. These results suggest that differences in distribution of 5-HT input may contribute to regional differences in PI muscle activation during inspiration and that differences in PI motoneuron recruitment do not relate to size.

Interaction between left and right intercostal muscles in airway pressure generation.

The interactions between the different rib cage inspiratory muscles in the generation of pleural pressure remain largely unknown. In the present study, we have assessed in dogs the interactions between the parasternal intercostals and the interosseous intercostals situated on the right and left sides of the sternum. For each set of muscles, the changes in airway opening pressure (DeltaPao) obtained during separate right and left activation were added, and the calculated values (predicted DeltaPao) were then compared with the DeltaPao values obtained during symmetric, bilateral activation (measured DeltaPao). When the parasternal intercostals in one or two interspaces were activated, the measured DeltaPao was commonly greater than the predicted value. The difference, however, was only 10%. When the interosseous intercostals were activated, the measured DeltaPao was nearly equal to the predicted value. These observations strengthen our previous conclusion that the pressure changes produced by the rib cage inspiratory muscles are essentially additive. As a corollary, the rib cage can be considered as a linear elastic structure over a wide range of distortion.

Abdominal muscle strength in patients with tetraplegia.

The abdominal muscles are completely paralyzed after traumatic transection of the cervical cord. To assess the residual pressure-generating capacity of these muscles, we first measured the changes in gastric pressure (DeltaPga) during paired bilateral stimulation of the lower thoracic nerve roots in eight chronic patients with C5-C7 tetraplegia and eight matched unaffected subjects in the seated posture. Stimulations were applied with a 90-mm circular magnetic coil positioned at the level of T10 and connected to a Magstim 250 stimulator. During relaxation at FRC, DeltaPga during maximal stimulation averaged (mean +/- SE) 76.0 +/- 11.7 cm H(2)O in the control subjects, whereas in the patients it was only 29.9 +/- 3.7 cm H(2)O (p = 0.002). Stimulations were next applied during the course of a forced expiration. All patients consistently demonstrated an abrupt increase in esophageal pressure (22.7 +/- 4.5 cm H(2)O), and six of them also showed an increase in expiratory flow. The cumulative thickness of the four abdominal muscles, as measured with an ultrasound probe, was 34% smaller in the patients than in the control subjects and correlated positively with maximal DeltaPga. We conclude that in patients with tetraplegia, muscle atrophy causes a marked reduction in abdominal muscle strength. However, magnetic stimulation of the abdominal muscles elicits increases in intrathoracic pressure that are greater than those required to initiate dynamic airway compression; it might, therefore, improve the clearing of airway secretions.

The canine parasternal and external intercostal muscles drive the ribs differently.

1. In the dog, the elevation of the ribs during inspiration results from the combined actions of the parasternal and external intercostal muscles. In the present studies, the hypothesis was tested that co-ordinated activity among these two sets of muscles reduces the distortion of the rib cage. 2. During spontaneous inspiration before or after section of the phrenic nerves, the ribs moved cranially and outward in the same way as they did during passive inflation. However, whereas the sternum moved cranially during passive inflation, it was displaced caudally during spontaneous inspiration. 3. When the parasternal intercostal muscles were selectively denervated, both the sternum and the ribs moved cranially, but the rib outward displacement was markedly reduced. In contrast, when the external intercostals were excised and the parasternal intercostals were left intact, the sternum continued to move caudally and the outward displacement of the ribs was augmented relative to their cranial displacement. 4. These observations establish that the external intercostal muscles drive the ribs primarily in the cranial direction, whereas the parasternal intercostals drive the ribs both cranially and outward. They also indicate, in agreement with the hypothesis, that co-ordinated activity among these two sets of muscles displaces the ribs on their relaxation curve. 5. However, this co-ordinated activity also displaces the sternum caudally. Although this distortion requires an additional energy expenditure, it enhances the outward component of rib displacement which is more effective with respect to lung expansion.

Response of the canine inspiratory intercostal muscles to chest wall vibration.

High-frequency mechanical vibration of the rib cage reduces dyspnea, but the effect of this procedure on the respiratory muscles is largely unknown. In the present studies, we have initially assessed the electrical and mechanical response to vibration (40 Hz) of the canine parasternal and external intercostal muscles (third interspace) during hyperventilation-induced apnea. When the vibrator was applied to the segment investigated, prominent external intercostal activity was recorded in the seven animals studied, whereas low-amplitude parasternal intercostal activity was recorded in only four animals. Similarly, when the vibrator was applied to more rostral and more caudal interspaces, activity was recorded commonly from the external intercostal but only occasionally from the parasternal. The two muscles, however, showed similar changes in length. We next examined the response to vibration of the muscles in seven spontaneously breathing animals. Vibrating the rib cage during inspiration (in-phase) had no effect on parasternal intercostal inspiratory activity but induced a marked increase in neural drive to the external intercostals. For the animal group, peak external intercostal activity during the control, nonvibrated breaths averaged (mean +/- SE) 43.1 +/- 3.7% of the activity recorded during the vibrated breaths (p < 0.001). External intercostal activity during vibration also occurred earlier at the onset of inspiration and commonly carried on after the cessation of parasternal intercostal activity. Yet tidal volume was unchanged. Vibrating the rib cage during expiration (out-of-phase) did not elicit any parasternal or external intercostal activity in six animals. These observations thus indicate that the external intercostals, with their larger spindle density, are much more sensitive to chest wall vibration than the parasternal intercostals. They also suggest that the impact of this procedure on the mechanical behavior of the respiratory system is relatively small.

Effects of increased ventilatory drive on motor unit firing rates in human inspiratory muscles.

This study was designed to determine whether increased neural drive increases firing rates of inspiratory motoneurons uniformly in humans. The discharge of single motor units in the diaphragm, parasternal intercostal and scalene muscles was recorded with monopolar electrodes. Ventilation was increased threefold with an external dead space. The discharge of 516 motor units was sampled in four subjects. All but 4 units increased their discharge rate during inspiration with only 46 discharging tonically during expiration. With increased dead space, discharge frequencies of diaphragmatic motor units increased from 11.0 +/- 2.7 to 17.7 +/- 3.3 Hz (mean +/- SD; p < 0.001). However, firing rates increased for parasternal intercostals from 10.0 +/- 1.6 to only 11.9 +/- 1.9 Hz (p < 0.001), and for scalenes from 8.7 +/- 1.8 to only 9.5 +/- 1.2 Hz (p < 0.05). Proportionate increases in rib cage and abdominal expansion accompanied the increased ventilation with added dead space. These results suggest that previously reported predominant increase in firing rates of diaphragmatic motor units in patients with chronic airflow limitation reflects the normal response of respiratory motor output to increased neural drive. The motoneuron pools of the parasternal intercostals and scalenes may show more prominent recruitment than frequency modulation.

Postinspiratory activity of the parasternal and external intercostal muscles in awake canines.

Previous studies have shown in awake dogs that activity in the crural diaphragm, but not in the costal diaphragm, usually persists after the end of inspiratory airflow. It has been suggested that this difference in postinspiratory activity results from greater muscle spindle content in the crural diaphragm. To evaluate the relationship between muscle spindles and postinspiratory activity, we have studied the pattern of activation of the parasternal and external intercostal muscles in the second to fourth interspaces in eight chronically implanted animals. Recordings were made on 2 or 3 successive days with the animals breathing quietly in the lateral decubitus position. The two muscles discharged in phase with inspiration, but parasternal intercostal activity usually terminated with the cessation of inspiratory flow, whereas external intercostal activity persisted for 24.7 +/- 12.3% of inspiratory time (P < 0.05). Forelimb elevation in six animals did not affect postinspiratory activity in the parasternal but prolonged postinspiratory activity in the external intercostal to 45.4 +/- 16.3% of inspiratory time (P < 0.05); in two animals, activity was still present at the onset of the next inspiratory burst. These observations support the concept that muscle spindles are an important determinant of postinspiratory activity. The absence of such activity in the parasternal intercostals and costal diaphragm also suggests that the mechanical impact of postinspiratory activity on the respiratory system is smaller than conventionally thought.

Muscle kinematics for minimal work of breathing.

A mathematical model was analyzed to obtain a quantitative and testable representation of the long-standing hypothesis that the respiratory muscles drive the chest wall along the trajectory for which the work of breathing is minimal. The respiratory system was modeled as a linear elastic system that can be expanded either by pressure applied at the airway opening (passive inflation) or by active forces in respiratory muscles (active inflation). The work of active expansion was calculated, and the distribution of muscle forces that produces a given lung expansion with minimal work was computed. The calculated expression for muscle force is complicated, but the corresponding kinematics of muscle shortening is simple: active inspiratory muscles shorten more during active inflation than during passive inflation, and the ratio of active to passive shortening is the same for all active muscles. In addition, the ratio of the minimal work done by respiratory muscles during active inflation to work required for passive inflation is the same as the ratio of active to passive muscle shortening. The minimal-work hypothesis was tested by measurement of the passive and active shortening of the internal intercostal muscles in the parasternal region of two interspaces in five supine anesthetized dogs. Fractional changes in muscle length were measured by sonomicrometry during passive inflation, during quiet breathing, and during forceful inspiratory efforts against a closed airway. Active muscle shortening during quiet breathing was, on average, 70% greater than passive shortening, but it was only weakly correlated with passive shortening. Active shortening inferred from the data for more forceful inspiratory efforts was approximately 40% greater than passive shortening and was highly correlated with passive shortening. These data support the hypothesis that, during forceful inspiratory efforts, muscle activation is coordinated so as to expand the chest wall with minimal work.

Spatial distribution of external and internal intercostal activity in dogs.

1. The observation that the external and internal interosseous intercostal muscles in the dog show marked regional differences in mechanical advantage has prompted us to re-examine the topographic distribution of electrical activity among these muscles during spontaneous breathing. 2. Inspiratory activity was recorded only from the areas of the external intercostals with an inspiratory mechanical advantage, and expiratory activity was recorded only from the areas of the internal intercostals with an expiratory mechanical advantage. The expiratory discharges previously recorded from the caudal external intercostals and the inspiratory discharges recorded from the rostral internal intercostals were probably due to cross-contamination. 3. Activity in each muscle area was also quantified relative to the activity measured during tetanic, supramaximal nerve stimulation (maximal activity). External intercostal inspiratory activity was consistently greater in the areas with a greater inspiratory advantage (i.e. the dorsal aspect of the rostral segments) than in the areas with a smaller inspiratory advantage, and internal intercostal expiratory activity was invariably greatest in the areas with the greatest expiratory advantage (i.e. the dorsal aspect of the caudal segments). 4. This topographic distribution of neural drive confers to the external intercostal muscles an inspiratory action on the lung during breathing and to the internal interosseous intercostals an expiratory action.

Respiratory mechanical advantage of the canine external and internal intercostal muscles.

1. The current conventional view of intercostal muscle actions is based on the theory of Hamberger (1749) and maintains that as a result of the orientation of the muscle fibres, the external intercostals have an inspiratory action on the lung and the internal interosseous intercostals have an expiratory action. This notion, however, remains unproved. 2. In the present studies, the respiratory actions of the canine external and internal intercostal muscles were evaluated by applying the Maxwell reciprocity theorem. Thus the effects of passive inflation on the changes in length of the muscles throughout the rib cage were assessed, and the distributions of muscle mass were determined. The fractional changes in muscle length during inflation were then multiplied by muscle mass and maximum active stress (3.0 kg cm-2) to evaluate the potential effects of the muscles on the lung. 3. The external intercostals in the dorsal third of the rostral interspaces were found to have a large inspiratory effect. However, this effect decreases rapidly both toward the costochondral junctions and toward the base of the rib cage. As a result, it is reversed to an expiratory effect in the most caudal interspaces. The internal intercostals in the caudal interspaces have a large expiratory effect, but this effect decreases ventrally and rostrally, such that it is reversed to an inspiratory effect in the most rostral interspaces. 4. These observations indicate that the canine external and internal intercostal muscles do not have distinct inspiratory and expiratory actions as conventionally thought. Therefore, their effects on the lung during breathing will be determined by the topographic distribution of neural drive.

Determinants of rib motion in flail chest.

We have previously developed a canine model of isolated flail chest to assess the effects of this condition on the mechanics of breathing, and these studies have led to the conclusion that the respiratory displacement of the fractured ribs is primarily determined by the fall in pleural pressure (Delta Ppl) and the action of the parasternal intercostal muscles. The present studies were designed to test the validity of this conclusion. A flail was induced in six supine anesthetized animals by fracturing both dorsally and ventrally the second to fifth ribs on the right side of the chest, after which the phrenic nerve roots were bilaterally sectioned in the neck. Sectioning the phrenic nerves caused a 34% decrease in Delta Ppl, associated with a 39% increase in parasternal intercostal inspiratory EMG activity (p < 0.05), and resulted in a marked reduction in the inspiratory inward displacement of the ribs. In three animals, the inward rib displacement was even reversed into a small outward displacement. When the airway was then occluded at end-expiration to increase Delta Ppl during the subsequent inspiration, all animals again showed a clear-cut inward rib displacement. These observations therefore confirm that in dogs with flail chest, the inspiratory displacement of the fractured ribs is set by the balance between the force related to pleural pressure and that generated by the parasternal intercostals. These observations also point to the critical importance of the pattern of inspiratory muscle activation in determining the magnitude of rib cage paradox in such patients.

Reflex inhibition of canine inspiratory intercostals by diaphragmatic tension receptors.

1. Electrical stimulation of phrenic afferent fibres in the dog elicits a reflex inhibition of efferent activity to the inspiratory intercostal muscles. However, electrical stimulation has a poor selectivity, so the sensory receptors responsible for this inhibition were not identified. 2. In the present studies, cranial forces were applied during spontaneous inspiration to the abdominal surface of the central, tendinous portion of the canine diaphragm to activate tension mechanoreceptors in the muscle. Vagal afferent inputs were eliminated by vagotomy. 3. The application of force to the central tendon caused a graded, reflex reduction in inspiratory intercostal activity, especially in external intercostal activity. This reduction was commonly associated with a decrease in inspiratory duration and was invariably attenuated after section of the cervical dorsal roots. 4. In contrast, no change in inspiratory intercostal activity was seen when high frequency mechanical vibration was applied to the central tendon to stimulate diaphragmatic muscle spindles. 5. These observations provide strong evidence that tension receptors in the diaphragm, but not muscle spindles, induce reflex inhibition of inspiratory intercostal activity. The expression of this reflex probably involves supraspinal structures.

Mechanical advantage of the human parasternal intercostal and triangularis sterni muscles.

1. Previous studies in dogs have demonstrated that the maximum change in airway pressure (DeltaPao) produced by a particular respiratory muscle is the product of three factors, namely the mass of the muscle, the maximal active muscle tension per unit cross-sectional area ( approximately 3.0 kg cm-2), and the fractional change in muscle length per unit volume increase of the relaxed chest wall (i.e. the muscle's mechanical advantage). In the present studies, we have used this principle to infer the DeltaPao values generated by the parasternal intercostal and triangularis sterni muscles in man. 2. The mass of the muscles and the direction of the muscle fibres relative to the sternum were first assessed in six cadavers. Seven healthy individuals were then placed in a computed tomographic scanner to determine the orientation of the costal cartilages relative to the sternum and their rotation during passive inflation to total lung capacity. The fractional changes in length of the muscles during inflation, their mechanical advantages, and their DeltaPao values were then calculated. 3. Passive inflation induced shortening of the parasternal intercostals in all interspaces and lengthening of the triangularis sterni. The fractional shortening of the parasternal intercostals decreased gradually from 7.7 % in the second interspace to 2.0 % in the fifth, whereas the fractional lengthening of the triangularis sterni increased progressively from 5.9 to 13.8 %. These rostrocaudal gradients were well accounted for by the more caudal orientation of the cartilages of the lower ribs. 4. Since these fractional changes in length corresponded to a maximal inflation, the inspiratory mechanical advantage of the parasternal intercostals was only 2.2-0. 6 % l-1, and the expiratory mechanical advantage of the triangularis sterni was only 1.6-3.8 % l-1. In addition, whatever the interspace, parasternal and triangularis muscle mass was 3-5 and 1-3 g, respectively. As a result, the magnitude of the DeltaPao values generated by a maximal contraction of the parasternal intercostals or triangularis sterni in all interspaces would be only 1-3 cmH2O. 5. These studies therefore confirm that the parasternal intercostals in man have an inspiratory action on the lung whereas the triangularis sterni has an expiratory action. However, these studies also establish the important fact that the pressure-generating ability of both muscles is substantially smaller than in the dog.