Three-dimensional reconstruction of human diaphragm with the use of spiral computed tomography

Pettiaux, Nicolas, Marie Cassart, Manuel Paiva, and MarcEstenne. Three-dimensional reconstruction of human diaphragm withthe use of spiral computed tomography. J. Appl.Physiol. 82(3): 998-1002, 1997.We developed atechnique of diaphragm imaging by using spiral computed tomography, andwe studied four normal subjects who had been previously investigatedwith magnetic resonance imaging (A. P. Gauthier, S. Verbanck,M. Estenne, C. Segebarth, P. T. Macklem, and M. Paiva.J. Appl. Physiol. 76: 495-506,1994). One acquisition of 15- to 25-s duration was performed atresidual volume, functional residual capacity, functional residualcapacity plus one-half inspiratory capacity, and total lung capacitywith the subject holding his breath and relaxing. From theseacquisitions, 20 coronal and 30 sagittal images were reconstructed ateach lung volume; on each image, diaphragm contour in the zone ofapposition and in the dome was digitized with the software Osiris, andthe digitized silhouettes were used for three-dimensionalreconstruction with Matlab. Values of length and surface area for thediaphragm, the dome, and the zone of apposition were very similar tothose obtained with magnetic resonance imaging. We conclude thatsatisfactory three-dimensional reconstruction of the in vivo diaphragmmay be obtained with spiral computed tomography, allowing accurate measurements of muscle length, surface area, and shape.

     

Diaphragm thickening during inspiration

Cohn, David, Joshua O. Benditt, Scott Eveloff, and F. DennisMcCool. Diaphragm thickening during inspiration.J. Appl. Physiol. 83(1): 291-296, 1997.Ultrasound has been used to measure diaphragm thickness(T_di) in thearea where the diaphragm abuts the rib cage (zone of apposition).However, the degree of diaphragm thickening during inspiration reportedas obtained by one-dimensional M-mode ultrasound was greater than thatpredicted by using other radiographic techniques. Becausetwo-dimensional (2-D) ultrasound provides greater anatomic definitionof the diaphragm and neighboring structures, we used this technique toreevaluate the relationship between lung volume andT_di. We firstestablished the accuracy and reproducibility of 2-D ultrasound bymeasuring T_diwith a 7.5-MHz transducer in 26 cadavers. We found thatT_di measured byultrasound correlated significantly with that measured by ruler (R² = 0.89), withthe slope of this relationship approximating a line of identity(y = 0.89x + 0.04 mm). The relationship between lung volume andT_di was thenstudied in nine subjects by obtaining diaphragm images at the fivetarget lung volumes [25% increments from residual volume (RV) tototal lung capacity (TLC)]. Plots ofT_di vs. lungvolume demonstrated that the diaphragm thickened as lung volumeincreased, with a more rapid rate of thickening at the higher lungvolumes[T_di = 1.74 vital capacity (VC)² + 0.26 VC + 2.7 mm] (R² = 0.99; P < 0.001) where lung volumeis expressed as a fraction of VC. The mean increase inT_di between RVand TLC for the group was 54% (range 42-78%). We conclude that2-D ultrasound can accurately measureT_di and that theaverage thickening of the diaphragm when a subject is inhaling from RVto TLC using this technique is in the range of what would be predictedfrom a 35% shortening of the diaphragm.

     

Effects of transverse fiber stiffness and central tendon on displacement and shape of a simple diaphragm model

Boriek, Aladin M., and Joseph R. Rodarte. Effects oftransverse fiber stiffness and central tendon on displacement and shapeof a simple diaphragm model. J. Appl. Physiol. 82(5): 1626-1636, 1997.Our previous experimental results (A. M. Boriek, S. Lui, and J. R. Rodarte. J. Appl. Physiol. 75:527-533, 1993 and A. M. Boriek, T. A. Wilson, and J. R. Rodarte.J. Appl. Physiol. 76: 223-229, 1994) showed that1) costal diaphragm shape is similar at functional residualcapacity and end inspiration regardless of whether the diaphragm muscleshortens actively (increased tension) or passively (decreased tension);2) diaphragmatic muscle length changes minimally in thedirection transverse to the muscle fibers, suggesting the diaphragm maybe inextensible in that direction; and 3) the central tendon isnot stretched by physiological stresses. A two-dimensional orthotropicmaterial has two different stiffnesses in orthogonal directions. In theplane tangent to the muscle surface, these directions are along thefibers and transverse to the fibers. We wondered whether orthotropicmaterial properties in the muscular region of the diaphragm andinextensibility of the central tendon might contribute to the constancyof diaphragm shape. Therefore, in the present study, we examined theeffects of stiffness transverse to muscle fibers and inextensibility ofthe central tendon on diaphragmatic displacement and shape. Finiteelement hemispherical models of the diaphragm were developed by usingpressurized isotropic and orthotropic membranes with a wide range ofstiffness ratios. We also tested heterogeneous models, in which themuscle sheet was an orthotropic material, having transverse fiberstiffness greater than that along the fibers, with the central tendonbeing an inextensible isotropic cap. These models revealed thatincreased transverse stiffness limits the shape change of thediaphragm. Furthermore, an inextensible cap simulating the centraltendon dramatically limits the change in shape as well as the membrane displacement in response to pressure. These findings provide a plausible mechanism by which the diaphragm maintains similar shapes despite different physiological loads. This study suggests that changesof diaphragm shape are restricted because the central tendon isessentially inextensible and stiffness in the direction transverse tothe muscle fibers is greater than stiffness along the fibers.

     

Muscle fiber architecture of the dog diaphragm

Boriek, Aladin M., Charles C. Miller III, and Joseph R. Rodarte. Muscle fiber architecture of the dog diaphragm.J. Appl. Physiol. 84(1): 318-326, 1998.Previous measurements of muscle thickness and length ratio ofcostal diaphragm insertions in the dog (A. M. Boriek and J. R. Rodarte.J. Appl. Physiol. 77: 2065-2070,1994) suggested, but did not prove, discontinuous muscle fiberarchitecture. We examined diaphragmatic muscle fiber architecture usingmorphological and histochemical methods. In 15 mongrel dogs, transversesections along the length of the muscle fibers were analyzedmorphometrically at ×20, by using the BioQuant System IVsoftware. We measured fiber diameters, cross-sectional fiber shapes,and cross-sectional area distributions of fibers. We also determinednumbers of muscle fibers per cross-sectional area and ratio ofconnective tissue to muscle fibers along a course of the muscle fromnear the chest wall (CW) to near the central tendon (CT) for midcostalleft and right hemidiaphragms, as well as ventral, middle, and dorsalregions of the left costal hemidiaphragm. In six other mongrel dogs,the macroscopic distribution of neuromuscular junctions (NMJ) onthoracic and abdominal diaphragm surfaces was determined by stainingthe intact diaphragmatic muscle for acetylcholinesterase activity. Theaverage major diameter of muscle fibers was significantly smaller, andthe number of fibers was significantly larger midspan between CT and CWthan near the insertions. The ratio of connective tissues to musclefibers was largest at CW compared with other regions along the lengthof the muscle. The diaphragm is transversely crossed by multiplescattered NMJ bands with fairly regular intervals offset in adjacentstrips. Muscle fascicles traverse two to five NMJ, consistent withfibers that do not span the entire fascicle from CT to CW. Theseresults suggest that the diaphragm has a discontinuous fiberarchitecture in which contractile forces may be transmitted among themuscle fibers through the connective tissue adjacent to the fibers.

     

Mechanisms for the mechanical response of airway smooth muscle to length oscillation

Shen, X., M. F. Wu, R. S. Tepper, and S. J. Gunst. Mechanisms for the mechanical response ofairway smooth muscle to length oscillation. J. Appl.Physiol. 83(3): 731-738, 1997.Airway smoothmuscle tone in vitro is profoundly affected by oscillations in musclelength, suggesting that the effects of lung volume changes on airwaytone result from direct effects of stretch on the airway smooth muscle.We analyzed the effect of length oscillation on active force andlength-force hysteresis in canine tracheal smooth muscle at differentoscillation rates and amplitudes during contraction with acetylcholine.During the shortening phase of the length oscillation cycle, the activeforce generated by the smooth muscle decreased markedly below theisometric force but returned to isometric force as the muscle waslengthened. Results indicate that at rates comparable to those duringtidal breathing, active shortening and yielding of contractile elementscontributes to the modulation of force during length oscillation;however, the depression of force during shortening cannot be accountedfor by cross-bridge properties, shortening-induced cross-bridgedeactivation, or active relaxation. We conclude that the depression ofcontractility may be a function of the plasticity of the cellularorganization of contractile filaments, which enables contractileelement length to be reset in relation to smooth muscle cell length asa result of smooth muscle stretch.

     

Instantaneous force-velocity-length relationship in diaphragmatic sarcomere

Coirault, Catherine, Denis Chemla, Jean-Claude Pourny,Francine Lambert, and Yves Lecarpentier. Instantaneousforce-velocity-length relationship in diaphragmatic sarcomere.J. Appl. Physiol. 82(2): 404-412, 1997.The simultaneous analysis of muscle force, length, velocity, andtime has been shown to precisely characterize the mechanicalperformance of isolated striated muscle. We tested the hypothesis thatthe three-dimensional force-velocity-length relationship reflectsmechanical properties of sarcomeres. In hamster diaphragm strips,instantaneous sarcomere length (SL) and muscle length were simultaneously measured during afterloaded twitches. SL was measured by means of laser diffraction. Wealso studied the influence of initialSL, abrupt changes in total load, and2 × 10⁷ M dantrolene.Baseline resting SL at the apex of thelength-active tension curve was 2.2 ± 0.1 µm, whereasSL at peak shortening was 1.6 ± 0.1 µm in the preloaded twitch and 2.1 ± 0.1 µm in the "isometric" twitch. Over the whole load continuum and at anygiven level of isotonic load, there was a unique relationship between instantaneous sarcomere velocity and instantaneousSL. Part of this relationship was timeindependent and initial SL independent and was markedly downshifted after dantrolene. When five different muscle regions were considered, there were no significant variations ofSL and sarcomere kinetics along themuscle. These results indicate that the time- and initiallength-independent part of the instantaneous force-velocity-lengthrelationship previously described in muscle strips reflects intrinsicsarcomere mechanical properties.

     

Diaphragm interference pattern EMG and compound muscle action potentials: effects of chest wall configuration

Beck, Jennifer, Christer Sinderby, Lars Lindström, andAlex Grassino. Diaphragm interference pattern EMG and compound muscle action potentials: effects of chest wall configuration. J. Appl. Physiol. 82(2): 520-530, 1997.The effect of chest wall configuration on the diaphragmelectromyogram (EMGdi) was evaluated in five healthy subjects with anesophageal electrode for both interference pattern EMGdi (voluntarycontractions) and electrically evoked diaphragm compound muscle actionpotentials (CMAPs). Diaphragm CMAPs (both unilateral and bilateral)were evaluated for the baseline-to-peak amplitude (Ampl), the time fromthe onset of the CMAP to first peak (T1), root mean square (RMS), andcenter frequency (CF) values of the CMAP power spectrum. CF values fromthe interference pattern EMGdi power spectrum were also calculated. ForCMAPs obtained at an electrode position least influenced by variationsinduced by electrode positioning, Ampl increased with diaphragmshortening from functional residual capacity (FRC) to total lungcapacity (TLC) by 101 and 98% (unilateral and bilateral,respectively). Bilateral CMAP RMS values increased 116% from FRC toTLC. CMAP T1 values decreased with diaphragm shortening from FRC to TLC by 1.1 and 2.1 ms for the unilateral and bilateral stimulations, respectively, and CF increased for the bilateral diaphragm CMAPs withdiaphragm shortening. CF values from the interference pattern EMGdi didnot show any consistent change with chest wall configuration. Thus CFvalues of the interference pattern EMGdi obtained with an esophagealelectrode can be considered reliable for physiological interpretation,at any diaphragm length (if electrode positioning and signalcontamination are controlled for), contrary to the diaphragm CMAPs,which are sensitive to changes in chest wall configuration. It isspeculated that the different results (over the effects of chest wallconfiguration on interference pattern EMGdi and diaphragm CMAPs) may bebecause of summation properties of the signals and how these influencethe EMG power spectrum.

     

Long-term facilitation of upper airway muscle activities in vagotomized and vagally intact cats

Mateika, J. H., and R. F. Fregosi. Long-termfacilitation of upper airway muscle activities in vagotomized andvagally intact cats. J. Appl. Physiol.82(2): 419-425, 1997.The primary purpose of the presentinvestigation was to determine whether long-term facilitation (LTF) ofupper airway muscle activities occurs in vagotomized and vagally intactcats. Tidal volume and diaphragm, genioglossus, and nasal dilatormuscle activities were recorded before, during, and after one carotidsinus nerve was stimulated five times with 2-min trains of constantcurrent. Sixty minutes after stimulation, nasal dilator andgenioglossus muscle activities were significantly greater than controlin the vagotomized cats but not in the vagally intact cats. Tidalvolume recorded from the vagotomized and vagally intact cats wassignificantly greater than control during the poststimulation period.In contrast, diaphragm activities were not significantly elevated inthe poststimulation period in either group of animals. We conclude that1) LTF of genioglossus and nasaldilator muscle activities can be evoked in vagotomized cats;2) vagal mechanisms inhibit LTF inupper airway muscles; and 3) LTF canbe evoked in accessory inspiratory muscles because LTF of inspiredtidal volume was greater than LTF of diaphragm activity.

     

Lung parenchymal shear modulus, airway wall remodeling, and bronchial hyperresponsiveness

Lambert, Rodney K., and Peter D. Paré. Lungparenchymal shear modulus, airway wall remodeling, and bronchialhyperresponsiveness. J. Appl. Physiol.83(1): 140-147, 1997.When airways narrow, either through theaction of smooth muscle shortening or during forced expiration, thelung parenchyma is locally distorted and provides an increasedperibronchial stress that resists the narrowing. Although thisinterdependence has been well studied, the quantitative significance ofairway remodeling to interdependence has not been elucidated. We haveused an improved computational model of the bronchial response tosmooth muscle agonists to investigate the relationships between airwaynarrowing (as indicated by airway resistance), parenchymal shearmodulus, adventitial thickening, and inner wall thickening at lungrecoil pressures of 4, 5, and 8 cmH₂O. We have found that, at lowrecoil pressures, decreases in parenchymal shear modulus have asignificant effect that is comparable to that of moderate thickening ofthe airway wall. At higher lung recoil pressures, the effect isnegligible.

     

Zone of apposition in the passive diaphragm of the dog

Boriek, Aladin M., Joseph R. Rodarte, and Susan S. Margulies. Zone of apposition in the passive diaphragm of thedog. J. Appl. Physiol. 81(5): 1929-1940, 1996.Wedetermined the regional area of the diaphragmatic zone of apposition(ZAP) as well as the regional craniocaudal extent of the ZAP(ZAP_ht) of the passive diaphragm in six paralyzedanesthetized beagle dogs (8-12 kg) at residual lung volume (RV),functional residual capacity (FRC), FRC + 0.25 and FRC + 0.5 inspiratory capacity, and total lung capacity (TLC) in prone and supinepostures. To identify the caudal boundary of the ZAP, 17 lead markers(1 mm) were sutured to the abdominal side of the costal and cruraldiaphragms around the diaphragm insertion on the chest wall. Two weekslater, the dogs' caudal thoraces were scanned by the use of thedynamic spatial reconstructor (DSR), a prototype fast volumetric X-raycomputer tomographic scanner, developed at the Mayo Clinic. Thethree-dimensional spatial coordinates of the markers were identified(±1.4 mm), and the cranial boundary of the ZAP was determined from30-40 1.4-mm-thick sagittal and coronal slices in each DSR image.We interpolated the DSR data to find the position of the cranial andcaudal boundaries of the ZAP every 5° around the thorax and computedthe distribution of regional variation of area of the ZAP andZAP_ht as well as the total area of ZAP. TheZAP_ht and area of ZAP increased as lung volume decreasedand were largest near the lateral extremes of the rib cage. We measuredthe surface area of the rib cage cephaled to the ZAP(A_L) in both postures in another six beagle dogs(12-16 kg) of similar stature, scanned previously in the DSR. Weestimated the entire rib cage surface area(A_rc = A_ZAP +A_L). The A_ZAP as a percentageof A_rc increased more than threefold as lung volumedecreased from TLC to RV, from ~9 to 29% of A_rc.

     

Effect of pulmonary emphysema on diaphragm capillary geometry

Poole, David C., and Odile Mathieu-Costello. Effect ofpulmonary emphysema on diaphragm capillary geometry.J. Appl. Physiol. 82(2): 599-606, 1997.In emphysema, the diaphragm shortens by losing sarcomeres. Wehypothesized that unless capillaries undergo a similar shortening,capillary geometry must be altered. Without quantifying this geometry,capillary length and surface area per fiber volume, which are criticalmeasurements of the structural potential for blood-tissue exchange,cannot be resolved. Five months after intratracheal elastase (E) orsaline (control; C) instillation, diaphragms from male Syrian goldenhamsters were glutaraldehyde perfusion fixed in situ at reference lungpositions (residual volume, functional residual capacity, total lungcapacity) to provide diaphragms fixed over a range of sarcomerelengths. Subsequently, diaphragms were processed for electronmicroscopy and analyzed morphometrically. Emphysema increased lungvolume changes from 20 to 25 cmH₂O airway pressure (i.e.,passive vital capacity) and excised lung volume (bothP < 0.001). In each region of thecostal diaphragm (i.e., ventral, medial, dorsal), sarcomere number wasreduced (all P < 0.05).Capillary-to-fiber ratio increased (C = 2.2 ± 0.1, E = 2.8 ± 0.1; P < 0.01) and fibershypertrophied (C = 815 ± 35, E = 987 ± 67 µm²;P < 0.05; both values at 2.5 µmsarcomere length). Capillary geometry was markedly altered by the lossof sarcomeres in series. Specifically, the additional capillary lengthderived from capillary tortuosity and branching was increased by 183%at 2.5 µm sarcomere length compared with C values (C, 359 ± 43;E, 1,020 ± 158 mm²,P < 0.01). This significantlyincreased total capillary length (C, 3,115 ± 173; E, 3,851 ± 219 mm² at 2.5 µm,P < 0.05) and surface area (C, 456 ± 13; E, 519 ± 24 cm¹,P < 0.05) per fiber volume. Thusemphysema substantially alters diaphragm capillary geometry andaugments the capillary length and surface area available forblood-tissue exchange.

     

Kinematics and mechanics of midcostal diaphragm of dog

Boriek, Aladin M., Joseph R. Rodarte, and Theodore A. Wilson. Kinematics and mechanics of midcostal diaphragm of dog. J. Appl. Physiol. 83(4):1068-1075, 1997.Radiopaque markers were attached to theperitoneal surface of three neighboring muscle bundles in the midcostaldiaphragm of four dogs, and the locations of the markers were trackedby biplanar video fluoroscopy during quiet spontaneous breathing andduring inspiratory efforts against an occluded airway at three lungvolumes from functional residual capacity to total lung capacity inboth the prone and supine postures. Length and curvature of the musclebundles were determined from the data on marker location. Musclelengths for the inspiratory states, as a fraction of length atfunctional residual capacity, ranged from 0.89 ± 0.04 at endinspiration during spontaneous breathing down to 0.68 ± 0.07 duringinspiratory efforts at total lung capacity. The muscle bundles werefound to have the shape of circular arcs, with the three bundlesforming a section of a right circular cylinder. With increasing lungvolume and diaphragm displacement, the circular arcs rotate around theline of insertion on the chest wall, the arcs shorten, but the radiusof curvature remains nearly constant. Maximal transdiaphragmaticpressure was calculated from muscle curvature and maximaltension-length data from the literature. The calculated maximaltransdiaphragmatic pressure-length curve agrees well with the data ofRoad et al. (J. Appl. Physiol. 60:63-67, 1986).

     

Stretch-induced enhancement of mechanical work production in frog single fibers and human muscle

Takarada, Yudai, Hiroyuki Iwamoto, Haruo Sugi, YuichiHirano, and Naokata Ishii. Stretch-induced enhancement ofmechanical work production in frog single fibers and human muscle.J. Appl. Physiol. 83(5):1741-1748, 1997.The relations between the velocity of prestretchand the mechanical energy liberated during the subsequent isovelocityrelease were studied in contractions of frog single fibers and humanmuscles. During isometric contractions of frog single fibers, a rampstretch of varied velocity (amplitude, 0.02 fiber length; velocity,0.08-1.0 fiber length/s) followed by a release (amplitude, 0.02 fiber length; velocity, 1.0 fiber length/s) was given, and the amountof work liberated during the release was measured. For human muscles,elbow flexions were performed with a prestretch of variedvelocity (range, 40°; velocity, 30-180°/s) followed by anisokinetic shortening (velocity, 90°/s). In both frog single fibersand human muscles, the work production increased with both the velocityof stretch and the peak of force attained before the release up to acertain level; thereafter it declined with the further increases ofthese variables. In human muscles, the enhancement of work productionwas not associated with a significant increase in integratedelectromyogram. This suggests that changes in intrinsic mechanicalproperties of muscle fibers play an important role in thestretch-induced enhancement of work production.

     

Pharmacological modulation of the mechanical response of airway smooth muscle to length oscillation

Shen, X., M. F. Wu, R. S. Tepper, and S. J. Gunst. Pharmacological modulation of the mechanicalresponse of airway smooth muscle to length oscillation.J. Appl. Physiol. 83(3): 739-745, 1997.Stretch and retraction of the airways caused by changes in lungvolume may play an important role in regulating airway reactivity. Westudied the effects of different pharmacological stimuli on airwaysmooth muscle to determine whether the muscle behavior during lengthoscillation can be modulated pharmacologically and to evaluate the roleof different activation mechanisms in determining its behavior duringthe oscillation. Active force decreased below the static isometricforce during the shortening phase of length oscillation, resulting inan overall depression of force during the length oscillation cycle.This pattern of response was unaffected by the contractile stimulus orlevel of activation, suggesting that it was caused by a mechanism that is independent of the level of activation of cross bridges. The normalized area of the length-force hysteresis loop (hysteresivity) differed depending on the stimulus used for contraction. Effects ofdifferent stimuli on hysteresivity were not correlated with theireffects on isotonic shortening velocity or isometric force, suggestingthat the pharmacological modulation of the behavior of airway smoothmuscle during length oscillation at these amplitudes cannot beaccounted for by the effects on the cross-bridge cycling rate.

     

Expression of the inducible nitric oxide synthase gene in diaphragm and skeletal muscle

Thompson, Marita, Lisa Becker, Debbie Bryant, Gary Williams,Daniel Levin, Linda Margraf, and Brett P. Giroir. Expression ofthe inducible nitric oxide synthase gene in diaphragm and skeletal muscle. J. Appl. Physiol. 81(6):2415-2420, 1996.Nitric oxide (NO) is a pluripotent molecule thatcan be secreted by skeletal muscle through the activity of the neuronalconstitutive isoform of NO synthase. To determine whether skeletalmuscle and diaphragm might also express the macrophage-inducible formof NO synthase (iNOS) during provocative states, we examined tissuefrom mice at serial times after intravenous administration ofEscherichia coli endotoxin. In thesestudies, iNOS mRNA was strongly expressed in the diaphragm and skeletalmuscle of mice 4 h after intravenous endotoxin and was significantlydiminished by 8 h after challenge. Induction of iNOS mRNA was followedby expression of iNOS immunoreactive protein on Western immunoblots.Increased iNOS activity was demonstrated by conversion of arginine tocitrulline. Immunochemical analysis of diaphragmatic explants exposedto endotoxin in vitro revealed specific iNOS staining in myocytes, inaddition to macrophages and endothelium. These results may be importantin understanding the pathogenesis of respiratory pump failure duringseptic shock, as well as skeletal muscle injury during inflammation ormetabolic stress.

     

Shape and size of the human diaphragm in vivo

Serial computerized tomograph (CT) sections at 5-mm intervals of a human diaphragm in relaxed and contracted states were obtained in one subject while he held his breath and lay supine in a CT scanner. All sections for one state were scanned at the same chest wall configuration as monitored by rib cage and abdominal dimensions, using magnetometers. Sections were scanned at relaxed functional residual capacity and after inspiring approximately 1 liter in such a way that rib cage dimensions increased only slightly. Models of the diaphragm dome in the two states were constructed from the sets of serial sections. Diaphragm length and volume displaced were measured, the zone of apposition of diaphragm to rib cage was mapped, and the line of the diaphragm silhouette in anteroposterior and lateral X-rays identified. Coronal and sagittal sections were constructed. In the inspiration studied, the diaphragm movement displaced 680 ml. Meridian lines in sagittal, coronal, and transverse directions over the right hemidiaphragm dome shortened by 6.7-7.2 cm, but over the left dome by only 4.0-4.3 cm. Lines of X-ray silhouettes were close to meridian lines, and estimates of shortening were similar to those made previously from X-rays. The peculiar saddle shape of the muscle may help the hemidiaphragms to operate independently, the fibers of the saddle acting as an anchor for midline directed fibers of the hemidiaphragm domes. The shape of the diaphragm also has implications for the distribution of transdiaphragmatic pressure and for the kind of distortion of the lower rib cage margin that is seen during inspirations at high lung volume.     

Hypothyroidism alters diaphragm muscle development

Sieck, Gary C., Louise E. Wilson, Bruce D. Johnson, andWen-Zhi Zhan. Hypothyroidism alters diaphragm muscle development. J. Appl. Physiol. 81(5):1965-1972, 1996.The impact of hypothyroidism (Hyp) onmyosin heavy chain (MHC) isoform expression, maximum specific force(P_o), fatigability, and maximumunloaded shortening velocity(V_o) wasdetermined in the rat diaphragm muscle (Dia) at 0, 7, 14, 21, and 28 days of age. Hyp was induced by treating pregnant rats with6-n-propyl-2-thiouracil (0.05% indrinking water) beginning at gestational day10 and was confirmed by reduced plasma levels of3,5,3-triiodothyronine and thyroxine. MHC isoforms wereseparated on sodium dodecyl sulfate-polyacrylamide gel electrophoresis gels and analyzed by densitometry. IsometricP_o and fatigue resistance of theDia were measured in vitro at 26°C, andV_o was determined at 15°C with the slack test. Compared with control muscles,expression of MHC-slow was higher and expression of adult fast MHCisoforms was lower in Hyp Dia at all ages. The neonatal isoform of MHC continued to be expressed in the Hyp Dia until day28. At each age,P_o and fatigability were reducedand V_o was slowerin the Hyp Dia. We conclude that Hyp-induced alterations in MHC isoform expression do not fully predict the changes in Dia contractile properties.

     

Sprint training, in vitro and in vivo muscle function, and myosin heavy chain expression

Harridge, S. D. R., R. Bottinelli, M. Canepari, M. Pellegrino, C. Reggiani, M. Esbjörnsson, P. D. Balsom, and B. Saltin. Sprint training, in vitro and in vivo muscle function, and myosin heavy chain expression. J. Appl.Physiol. 84(2): 442-449, 1998.Sprint trainingrepresents the condition in which increases in muscle shortening speed,as well as in strength, might play a significant role in improvingpower generation. This study therefore aimed to determine the effectsof sprint training on 1) thecoupling between myosin heavy chain (MHC) isoform expression andfunction in single fibers, 2) thedistribution of MHC isoforms across a whole muscle, and3) in vivo muscle function. Sevenyoung male subjects completed 6 wk of training (3-s sprints) on a cycleergometer. Training was without effect on maximum shortening velocityin single fibers or in the relative distribution of MHC isoforms ineither the soleus or the vastus lateralis muscles. Electrically evokedand voluntary isometric torque generation increased(P < 0.05) after training in boththe plantar flexors (+8% at 50 Hz and +16% maximal voluntarycontraction) and knee extensors (+8% at 50 Hz and +7% maximalvoluntary contraction). With the shortening potential of the musclesapparently unchanged, the increased strength of the major lower limbmuscles is likely to have contributed to the 7% increase(P < 0.05) in peak pedal frequency during cycling.

     

Enhancement of signal quality in esophageal recordings of diaphragm EMG

Sinderby, Christer A., Jennifer C. Beck, Lars H. Lindström, and Alejandro E. Grassino. Enhancement of signalquality in esophageal recordings of diaphragm EMG. J. Appl. Physiol. 82(4): 1370-1377, 1997.The cruraldiaphragm electromyogram (EMGdi) is recorded from a sheet of muscle,the fiber direction of which is mostly perpendicular to an esophagealbipolar electrode. The region from which the action potentials areelicited, the electrically active region of the diaphragm(EAR_di) and the center of this region (EAR_{di ctr}) mayvary during voluntary contractions in terms of their position withrespect to an esophageal electrode. Depending on the bipolarelectrode's position with respect to theEAR_{di ctr}, the EMGdi isfiltered to different degrees. The objectives of the present study wereto reduce these filtering effects on the EMGdi by developing ananalysis algorithm referred to as the "double-subtraction technique." The results showed that changes in the position of theEAR_{di ctr} by ±5 mm withrespect to the electrode pairs located 10 mm caudal and 10 mm cephaladprovided a systematic variation in the EMG power spectrumcenter-frequency values by ±10%. The double-subtraction techniquereduced the influence of movement of theEAR_{di ctr} relative to theelectrode array on EMG power spectrum center frequency and root meansquare values, increased the signal-to-noise ratio by 2 dB, andincreased the number of EMG samples that were accepted by the signalquality indexes by 50%.

     

Functional role and structure of the scalene: an accessory inspiratory muscle in hamster

Fournier, Mario, and Michael I. Lewis. Functional roleand structure of the scalene: an accessory inspiratory muscle inhamster. J. Appl. Physiol. 81(6):2436-2444, 1996.Although the scalene muscle (Sca) is a primaryinspiratory muscle in humans, its respiratory function in other speciesis less clear. The electromyographic (EMG) activity of the Sca wasstudied during resting ventilation (eupnea) in both the awake andanesthetized hamster and after a variety of respiratory challenges inthe anesthetized animal. The EMG activities of the medial Sca and thecostal diaphragm were compared. The medial Sca, the major component ofthe Sca, originates from cervical transverse processes 2 to 5 andinserts primarily onto rib 4, with a small segment onto rib 3. In both the anesthetized and awake animal, the Sca was always silent during quiet breathing. WithCO₂-stimulated hyperpnea, the Scawas always recruited during inspiration in phase with the diaphragm.Active recruitment of the Sca was also observed after resistive loading and total airway occlusion. After ipsilateral phrenicotomy, the Sca waspersistently recruited during eupnea. The specificity of the EMGsignals was tested both by excluding cross contamination from other ribcage muscles and by selective denervation studies. Muscle spindles wereidentified in the medial Sca histochemically, suggesting that therespiratory activity of the Sca can also be modulated by changes inmuscle length and/or load. These results indicate that the Scafunctions as an accessory inspiratory muscle in the hamster and mayplay an important role in conditions of chronic load.