Internal Intercostal Research Articles

EMG recordings of the respiratory muscles during unilateral and bilateral chest expansion

Respiratory muscle (RM) EMG was recorded in 10 healthy adults during bilateral and voluntary unilateral inspiration. Each performed bilateral and unilateral inspiration at 50, 75 and 100 per cent of maximal inspiration (PI max) measured with a mouth pressure gauge. The obliquus externus abdominus (EAO) EMG was also recorded in five subjects. Results showed that RM and EAO activity on the expanding side was similar during all levels of effort. The RM activity on the non-contracting side during most submaximal efforts was higher than that on the same side during bilateral efforts but not during maximal effort. The increase in EMG could be due to cross-talk from EAO and latissimus dorsi, or co-activation of the internal intercostal muscles. Because RM activity at PI max was similar for unilateral and bilateral efforts, it was concluded that voluntary unilateral RM inhibition may not be possible during maximal effort.

Expiratory activity of the inspiratory muscles during cough.

To investigate the neural mechanism of the expiratory activity of the inspiratory muscles during a cough, EMG of the respiratory muscles were recorded in anesthetized and tracheostomized dogs. A laparoscope was used to minimize injury to the abdominal muscles for implantation of the electrodes into the costal diaphragm. During the expulsive phase of a cough, the diaphragm was active in 7 of 12 dogs and the external intercostal muscle was active in 3 of 6 dogs. During a cough, the expiratory activity of the diaphragm, after the termination of its inspiratory activity, started at 52.9 +/- 24.6 ms, and that of external intercostal muscle started at 51.1 +/- 20.5 ms. The expiratory activity of the internal intercostal muscle and of the transversus abdominis started at 34.3 +/- 13.0 and 27.8 +/- 15.2 ms, respectively. The onset of expiratory activity of the inspiratory muscles is significantly later than that of expiratory muscles. Continuous activity in the expiratory muscles evoked by airway occlusion, i.e., Hering-Breuer reflex, was suppressed during the inspiratory phase of a cough, but not suppressed during the expulsive phase even when the expiratory activity of the diaphragm was observed. We concluded that the expiratory activity of inspiratory muscles is controlled independently of both expiratory activity of the expiratory muscles and inspiratory activity of the inspiratory muscles.

Inhomogeneous response of expiratory muscle activity to cold block of the ventral medullary surface

We assessed the effects of cooling the ventral medullary surface (VMS) on the activity of chest wall and abdominal expiratory muscles in eight anesthetized artificially ventilated dogs after vagotomy and denervation of the carotid sinus nerves. Electromyograms (EMGs) of the triangularis sterni, internal intercostal, abdominal external oblique, abdominal internal oblique, and transversus abdominis muscles were measured with EMG of the diaphragm as an index of inspiratory activity. Bilateral localized cooling (2 x 2 mm) in the thermosensitive intermediate part of the VMS produced temperature-dependent reduction in the EMG of diaphragm and abdominal muscles. The rib cage expiratory EMGs were little affected at 25 degrees C; their amplitudes decreased at lower VMS temperatures (less than 20 degrees C) but by significantly fewer degrees than the diaphragmatic and abdominal expiratory EMGs at a constant VMS temperature. With moderate to severe cooling (less than 20 degrees C) diaphragmatic EMG disappeared, but rib cage expiratory EMGs became tonic and resumed a phasic pattern shortly before the recovery of diaphragmatic EMG during rewarming of the VMS. These results indicate that the effects of cooling the VMS differ between the activity of rib cage and abdominal expiratory muscles. This variability may be due to inhomogeneous inputs from the VMS to expiratory motoneurons or to a different responsiveness of various expiratory motoneurons to the same input either from the VMS or the inspiratory neurons.

Blood flow distribution within the rib cage muscles

We used 15-microns radiolabeled microspheres to study the regional distribution of blood flow (Q) among parasternal (PS), transversus thoracis, and external (EI) and internal intercostal muscles (II) in nine anesthetized supine mongrel dogs. We measured Q (ml.min-1.100 g-1) in each intercostal space (ICS) during spontaneous breathing, inspiratory resistive loading, and mechanical ventilation following paralysis. At necropsy the EI, II, and PS were excised and sampled separately for each ICS. During paralysis there was no consistent gradient in Q among the PS, II, and EI muscles. During spontaneous breathing, Q to PS increased linearly by 125% between the first and fourth to sixth ICS, Q to EI decreased progressively from the first/second ICS to the fifth/sixth ICS, whereas Q to the II was uniform. During inspiratory resistive loading, in which mouth pressures of -16 +/- 4 cmH2O were generated, the PS gradient was similar to that during spontaneous breathing. Also, Q to the EI increased in the cranial interspaces (P less than 0.02), whereas Q to the II of the seventh/eighth ICS was greater than that of the first/second ICS (P less than 0.001). Furthermore, with loading, ventrodorsal gradients in Q appeared within both EI and II interspaces. There was no consistent gradient in Q within the transversus thoracis muscle during any of the interventions. Our results demonstrate nonuniform Q within PS, EI, and II during both spontaneous and inspiratory resistive loaded breathing. On the assumption that changes in Q reflect changes in activation, our results suggest systematic topographical patterns of recruitment of rib cage respiratory muscles.

Intercostal muscle action inferred from finite-element analysis

The external and internal intercostal muscles are important respiratory muscles in humans, but their mechanical actions have been controversial. We used finite-element analysis based on anatomic and mechanical measurements in dogs to assess the action of the intercostal and other rib cage muscles in a model of an isolated canine rib cage. When intercostal muscle forces of either the internal or the external layer were applied in a single interspace, they pulled the adjacent ribs together, consistent with published observations in dogs. However, when the forces were applied in all interspaces, the external layer caused an inspiratory motion and the internal layer caused an expiratory motion, consistent with conventional understanding of intercostal muscle actions. Parasternal intercostal, levator costae, and transversus thoracis (triangularis sterni) muscle actions were also simulated. These muscles caused expected movements of the ribs and sternum. We conclude that the actions of intercostal muscles depend on the spatial extent of their activation. Their actions in a single interspace and in multiple interspaces can be observed and explained with three-dimensional finite-element models.

Action of intercostal muscles on the lung in dogs

The action on the lung of interosseous intercostal muscles located in the third and the seventh interspaces was studied in 15 anesthetized-curarized supine dogs. Changes in pleural pressure, airflow rate, and lung volume produced by maximal stimulation of both intercostal muscle layers were measured at and above functional residual capacity (FRC). In five animals measurements were also obtained during isolated stimulation of the internal layer. At FRC, intercostal stimulation in the upper interspaces had invariably an inspiratory effect on the lung but no effect was detectable in the lower interspaces. Qualitatively similar results were obtained during isolated stimulation of the internal layer. Increasing lung volume reduced the inspiratory action of the upper intercostals and conferred an expiratory action to the lower intercostals. These results indicate the following: 1) when contracting in a single interspace, the external and internal intercostals have a qualitatively similar action on the lung; and 2) this action, however, depends critically on their location along the cephalocaudal axis of the rib cage: in the upper portion of the rib cage, both muscle layers have an inspiratory effect at and above FRC; in the lower portion of the rib cage, they have no respiratory action at FRC and act in the expiratory direction at higher lung volumes.

Rostrocaudal variation of fiber type composition in rat intercostal muscles

We used the histochemical stain for ATPase to compare the fiber-type composition of rat internal and external intercostal muscles from thoracic (T) segments 2-5, 8, and 11. At each level, type II fibers were more numerous than type I fibers, type II B fibers were more numerous than II A fibers, and type I fibers were more numerous in external than in internal intercostals. However, fiber type composition varied from segment to segment. For example, the proportion of type II A fibers increased in a rostrocaudal gradient in internal but not external intercostals, and type I fibers were more prevalent at rostral and caudal than at intermediate levels in both internal and external intercostals. These results provide a basis for interpreting previous physiological and molecular studies which have compared intercostal muscles from different segmental levels.

Respiratory Muscle Activity during Vocalization in the Squirrel Monkey

In order to find out which muscles are involved in the respiratory component of primate phonation, the activity of 17 abdominal and thoracic muscles was recorded during vocalization in the squirrel monkey. Vocalization-correlated activity was found in the musculi obliquus externus et internus, rectus et transversus abdominis, intercostalis externus et internus and intercartilagineus. It was lacking in the mm. iliocostalis, latissimus dorsi, longissimus dorsi rhomboideus, serratus posterior superior, trapezius, splenius capitis, sternocleidomastoideus, scalenus medius and pectoralis major. There was simultaneous activation of the rib-raising external and rib-lowering internal intercostal muscles during most vocalizations. It is hence concluded that the intercostals, rather than supporting expiratory efforts, serve to stabilize the thorax, thus providing an anchorage against which the abdominal muscles can act.

Regional blood flow to canine parietal pleura and internal intercostal muscle

Transcapillary Starling forces in the parietal pleura and the underlying interstitium may potentially contribute to the exchange of fluid across this barrier. However, the extent of blood flow to the parietal pleura has not been measured. Thus, using standard microsphere techniques, we compared blood flow to the parietal pleura, including the subpleural interstitium, with blood flow to the adjacent internal intercostal muscle, as well as with flows to other serous tissues, including mediastinal pleura, pericardium, and parietal peritoneum, in anesthetized dogs that were either breathing spontaneously (n = 9) or ventilated to control arterial PCO2 (n = 5). Blood flow (ml.min-1.g-1) was measured after 20 min of equilibration in four successive body positions: right lateral decubitus, supine, left lateral decubitus, and prone. Overall, flow to parietal pleura was not different in spontaneous [1.07 +/- 0.14 (SE)] and mechanically ventilated animals (0.74 +/- 0.11). Flow to the internal intercostal muscle was significantly less than pleural blood flow, averaging 0.24 +/- 0.03 and 0.16 +/- 0.03 in the same groups, although again there was no effect of ventilation mode. Blood flow to other serous tissues in the thoracic cavity, specifically the mediastinal pleura (0.67 +/- 0.14) and pericardium (0.88 +/- 0.22), was similar to parietal pleural flow, whereas that to the parietal peritoneum was an order of magnitude lower (0.09 +/- 0.02, P less than 0.05). Changing body position had no effect on blood flow to any of the sampled tissues. Blood flow to the dorsal aspect of the chest wall muscle in spontaneously breathing animals tended to be greater than that to lateral or ventral portions of the chest wall.(ABSTRACT TRUNCATED AT 250 WORDS)

Influence of body posture and lung volume on subglottal pressure control during singing

Summary The role of different breathing muscles during singing was investigated by synchronously recording EMG, pressure, and sound signals, using lung volume and gravity as experimental parameters. Surface EMG signals from the external and internal intercostals, the diaphragm, and the abdominal oblique muscles were recorded, while two singer subjects performed various singing tasks associated with rapid and precise changes of subglottal pressure. Esophageal and gastric pressures were measured by pressure transducers, and lung volume by means of impedance plethysmography. The results show that the breathing system efficiently compensates for drastic differences in the mechanics of the breathing apparatus, caused by differences in lung volume and gravity induced by changes of body posture.

Influences of Lung Mechanoreceptors and Carotid Chemoreceptors on the Response of Respiratory Muscle Activity to Tracheal Occlusion.

We examined the responses of respiratory muscle electromyograms (EMGs) from internal (IIC) and external intercostal (EIC) muscles and diaphragm (DIAP) to three successive occluded breaths in anesthetized spontaneously breathing rabbits. Both inspiratory and expiratory muscle EMGs progressively increased in the course of tracheal occlusion. An increase in these muscle EMGs was still observed after release of tracheal occlusion, but those effects were short-lasting. In a separate series of experiments, for assessment of possible reflex effects involved, the responses of slowly adapting pulmonary stretch receptor (SAR), rapidly adapting pulmonary stretch receptor (RAR), and carotid chemoreceptor activities to tracheal occlusion lasting for three respiratory efforts were also examined. The inspiratory discharge of SARs decreased but the expiratory discharge of SARs increased during tracheal occlusion. Although carotid chemoreceptors increased their activity in the latency of 3-6s after the onset of tracheal occlusion, the activity of RARs was greatly reduced throughout the period of tracheal occlusion. A transient increase in both carotid chemoreceptors and RARs was still observed after release of tracheal occlusion. These results suggest that alterations of inspiratory and expiratory muscle EMGs produced by tracheal occlusion would appear to be mediated by the afferent inputs from lung mechanoreceptors and carotid chemoreceptors.

Mechanical Action of the Interosseous Intercostal Muscles as a Function of Lung Volume

On the basis of local stimulation of individual muscles, it has been suggested that both the external (EI) and internal interosseous intercostal muscles have an inspiratory action at low lung volumes and an expiratory action at high lung volumes. In this study, we assessed the action of the interosseous intercostal muscles at different lung volumes in 19 anesthetized dogs by synchronously activating the intercostal muscles via ventral root stimulation (VRS). An electrode was positioned on the upper thoracic spinal cord according to previously described techniques. The cervical phrenic rootlets were sectioned bilaterally, the accessory muscles were sectioned from the rib cage, and the origins of the abdominal muscles were sectioned from the middle region of the rib cage. Changes in airway pressure (delta P) were monitored during the application of supramaximal stimuli after hyperventilation-induced apnea and during airway occlusion. Animals were passively inflated or deflated with a volume syringe. Precontractile airway pressure was used as an index of lung volume. External and parasternal intercostal muscle (PA) lengths were monitored by sonomicrometry in the third intercostal space. Thoracoabdominal motion was monitored by Respitrace bands. During VRS, both PA and EI shortened at all lung volumes. Mean delta P progressively decreased with increasing lung volume. At precontractile airway pressures of -10, 0, and +30 cm H2O, delta P were -25 +/- 1, -16 +/- 1, and -5 +/- 1 cm H2O, respectively. After section of the internal intercostal nerves lateral to the costochondral junctions from the first through the seventh intercostal spaces to eliminate PA action, EI shortened, whereas PA usually lengthened.(ABSTRACT TRUNCATED AT 250 WORDS)

Distribution of muscle fiber types and EMG activity in cat intercostal muscles

The electromyogram (EMG) activity and histochemical properties of intercostal muscles in the anesthetized cat were studied. The parasternal muscles were consistently active during inspiration. The external intercostals in the rostral spaces and the ventral portions of the midthoracic spaces were also recruited during inspiration. The remaining external intercostals were typically silent, regardless of the level of respiratory drive. The internal intercostal muscles located in the caudal spaces were occasionally recruited during expiration. There was a clear correlation between recruitment patterns of the intercostals and the histochemically defined fiber type properties of the muscles. Intercostal muscles that were routinely recruited during inspiration had a significantly higher proportion of slow-oxidative muscle fibers.

After phrenicotomy the rat alters the output of the remaining respiratory muscles without changing its sleep-waking pattern

The respiratory activity of selected rib cage and abdominal wall muscles was studied in intact and bilaterally phrenicotomized (PNX) rats during non-rapid eye movement sleep (nREMS) and REMS. The polygraphic method was used to identify the animal's sleep-waking states before and after PNX. Electromyographic (EMG) recordings were made from the following muscles: the parasternals of the 1st, 3rd, and 5th interspaces; the external and internal intercostals of the 1st, 6th and 10th interspaces; the levator costae attaching to the 10th rib; the scalenus medius; and, the abdominal wall muscles, teh external and internal obliques and rectus abdominis. Afte PNX, all rib cage muscles contracted exclusively during inspiration and all but one increased their activity. The exception was the internal intercostal muscl of the 10th interspace; its activity decreased. The external and internal oblique muscles, both of which were active during expiration in nREMS, also increased their output after PNX: rectus abdominis became an inspiratory muscle. The persistence of phasic activity of respiratory muscles during REMS varied not only from muscle to muscle but from one REMS epoch to another. The sleep-waking pattern of the PNX rat differed in only a minor way from that of the intact rat. Therefore, we conclude that the rat with total paralysis of its diaphragm uses mainly its neurometabolic mechanisms to achieve an adequate level of alveolar ventilation and not neurobehavioral mechanisms.

Respiratory Muscle Fiber Morphometry: Correlation with Pulmonary Function and Nutrition

To examine the relationship between nutrition, pulmonary function, respiratory muscle strength, and respiratory muscle morphometry, we compared physiologic data and muscle morphometry obtained from internal intercostal, external intercostal, and latissimus dorsi muscle biopsies in 68 patients who were having a thoracotomy. We stained the biopsies for myosin ATPase and measured the proportions and diameters of the type 1 and type 2 fibers. There were more qualitative changes in the external intercostal muscles than in the other two, and some of these changes related to the incidence of malignancy. There were more type 1 fibers in the external intercostal (64 +/- 10 percent) and internal intercostal muscles (59 +/- 12 percent) than in the latissimus dorsi (44 +/- 13 percent) (p less than 0.005). The mean diameter of the type 2 fibers in the external intercostal muscles was less (44 mu +/- 7 mu) than the diameter in the latissimus dorsi (51 mu +/- 9 mu) and the internal intercostal muscles (52 mu +/- 8 mu) (p less than 0.01). The diameters of both fiber types were greater in men than in women. There was no significant relationship between measures of pulmonary function or respiratory muscle strength and muscle fiber proportions and diameters. There were significant correlations between the percentage of ideal body weight and type 1 and type 2 fiber diameters. We conclude that sex and nutrition influence respiratory muscle morphometry.

Respiratory muscle control during vomiting

The changes in thoracic and abdominal pressure that generate vomiting are produced by coordinated action of the major respiratory muscles. During vomiting, the diaphragm and external intercostal (inspiratory) muscles co-contract with abdominal (expiratory) muscles in a series of bursts of activity that culminates in expulsion. Internal intercostal (expiratory) muscles contract out of phase with these muscles during retching and are inactive during expulsion. The periesophageal portion of the diaphragm relaxes during expulsion, presumably facilitating rostral movement of gastric contents. Recent studies have begun to examine to what extent medullary respiratory neurons are involved in the control of these muscles during vomiting. Bulbospinal expiratory neurons in the ventral respiratory group caudal to the obex discharge at the appropriate time during (fictive) vomiting to activate either abdominal or internal intercostal motoneurons. The pathways that drive phrenic and external intercostal motoneurons during vomiting have yet to be identified. Most bulbospinal inspiratory neurons in the dorsal and ventral respiratory groups do not have the appropriate response pattern to initiate activation of these motoneurons during (fictive) vomiting. Relaxation of the periesophageal diaphragm during vomiting could be brought about, at least in part, by reduced firing of bulbospinal inspiratory neurons.

Responses of inspiratory neurons of the dorsal respiratory group to stimulation of expiratory muscle and vagal afferents

In decerebrate, paralyzed and ventilated cats, we monitored the intracellular responses of 30 inspiratory neurons of the dorsal respiratory group (DRG) to stimulation of vagal and expiratory muscle (internal intercostal and abdominal) afferents. We hypothesized that the inhibitory effects of stimulation of expiratory muscle afferents, previously reported, would block the excitatory responses of inspiratory neurons of the DRG to vagal stimulation. Although prolonged stimulus trains to expiratory muscle afferents caused respiratory phase-switching, single shocks or short trains elicited no responses in 17 bulbospinal neurons, excitatory responses in 6, and inhibitory responses in 2. Of the 4 propriobulbar neurons tested, 2 had inhibitory responses and 2 did not respond. In only 2 neurons, both bulbospinal, did conditioning stimuli to expiratory muscle afferents block or reduce the excitatory effects of vagal stimulation. These results suggest that interaction of vagal and expiratory muscle afferents, which might account for the absence of a change in inspiratory duration despite increased vagal afferent feedback at elevated end-expiratory lung volumes, does not occur within the DRG.

Length changes of intercostal muscles during respiration in the cat

Sonomicrometry was used to measure the length changes of intercostal muscles thoughout the rib cage of anesthetized cats. The motor unit discharge in the vicinity of the length measurement was monitored with bipolar electrodes. The external intercostal muscles in the rostral spaces and the parasternals actively shortened during inspiration. The external intercostals in the caudal spaces, which were inactive, either shortened or lengthened passively due to the forces imposed on the lower rib cage by the contraction of the diaphragm. The length changes of external intercostals of the mid-thoracic spaces, which were recruited during inspiration, were variable. The internal intercostals were usually inactive in our preparation and therefore their length excursions passively followed those of the external intercostals. The information regarding the length changes is correlated with that previously gathered for the role of muscle spindles in the intercostal muscles. It is suggested that γ motoneurons are recruited to prevent the muscle spindles from being unloaded when the intercostal muscle shortens and to increase the sensitivity of the receptors during the phase of the respiratory cycle when the muscle is active.

Intercostal and abdominal muscle afferent influence on caudal medullary expiratory neurons that drive abdominal muscles

Our objective was to determine if caudal ventral respiratory group (VRG) expiratory (E) neurons that drive abdominal expiratory motoneurons in the lumbar cord respond to intercostal and lumbar nerve afferent stimulation. Results showed that 92% of medullary E-neurons that were antidromically activated from the upper lumbar cord reduced their activity in response to stimulation of external and internal intercostal and lumbar nerve afferents. We conclude that afferent information from intercostal and abdominal muscle tendon organs has an inhibitory effect on caudal VRG E-neurons that drive abdominal expiratory motoneurons.

Electrode for selective recording of electromyograms from intercostal muscles

An electromyogram-recording electrode is described that makes it possible to record separately the electrical signals generated within two closely approximated muscle layers. The device consists of two bipolar wire hook electrodes embedded in opposite faces of a thin laminated plastic wafer. The middle lamina of the wafer is a sheet of metal foil that shields the electrical field on one side of the wafer from the bipolar electrode on the other side. The device was tested by inserting it from the inside of the chest wall between the internal and external intercostal muscle layers. Signals from the two muscle layers were clearly separated. Single motor unit spikes were attenuated by factors ranging from 41 to 2.4. The device can be implanted with minimal trauma to surrounding muscles and is suitable for chronic animal experiments.