Mechanism of triangularis sterni shortening during expiration in dogs

The triangularis sterni in the dog shortens during expiration below its in situ relaxation length (Lr) (J. Appl. Physiol. 61: 539-544, 1986). To assess the mechanism of this expiratory muscle shortening, we have measured the electromyogram and the respiratory changes in length of the canine triangularis sterni in the third and fourth right intercostal space, first before and then after selective denervation. Eleven anesthetized, spontaneously breathing animals were studied in the supine posture; five of them were also studied during postural changes from supine to head-up. Before denervation, the muscle in the supine animals shortened by -12.84 +/- 1.81% Lr. With selective denervation, the amount of expiratory muscle shortening was reduced to only -2.54 +/- 0.71% Lr (P less than 0.001). Similarly, a change from the supine to the head-up posture before denervation promoted an increase in expiratory muscle shortening from -13.58 +/- 3.62 to -21.17 +/- 4.04% Lr (P less than 0.005), but the denervation abolished this increase. Denervating the triangularis sterni, however, did not affect expiratory activation of the internal intercostals. These results demonstrate that the expiratory contraction of the canine triangularis sterni is agonistic in nature, and they suggest that this contraction is responsible for most of the active caudal displacement of the ribs in the upper half of the rib cage.     

Coupling between triangularis sterni and parasternals during breathing in dogs

The purpose of the present studies was to assess the functional coupling between the parasternal intercostals and the triangularis sterni (transversus thoracis) muscles during resting breathing, and we measured the electrical activity and the respiratory changes in length of these two muscles in 13 supine anesthetized dogs. The changes in muscle length were defined relative to their respective in situ relaxation length (Lr). During inspiration, the parasternal intercostals were active and shortened below Lr, causing the triangularis sterni to be passively stretched above Lr. Shortly after the cessation of parasternal contraction, the triangularis sterni became active and shortened below Lr, and in nine animals this active shortening was associated with a forcible distension of the parasternal intercostals above Lr. Deactivation of the triangularis sterni at end expiration caused both muscles to return to their respective Lr. This pattern was essentially unchanged after supplemental anesthesia and bilateral phrenicotomy. We conclude that in dogs breathing quietly the length of the rib cage muscles during the expiratory pause is not passively determined as conventionally thought.     

Mechanical advantage of the canine triangularis sterni

De Troyer, André, and Alexandre Legrand.Mechanical advantage of the canine triangularis sterni.J. Appl. Physiol. 84(2): 562-568, 1998.Recent studies on the canine parasternal intercostal,sternomastoid, and scalene muscles have shown that the maximal changesin airway opening pressure (Pao) obtained per unit muscle mass(Pao/m) during isolatedcontraction are closely related to the fractional changes in musclelength per unit volume increase of the relaxed chest wall. In thepresent study, we have examined the validity of this relationship for the triangularis sterni, an important expiratory muscle of the rib cagein dogs. Passive inflation above functional residual capacity (FRC)induced a virtually linear increase in muscle length, such that, with a1.0-liter inflation, the muscle lengthened by 17.9 ± 1.6 (SE) % of its FRC length. When the muscle in one interspace wasmaximally stimulated at FRC, Pao increased by 0.84 ± 0.11 cmH₂O. However, in agreement withthe length-tension characteristics of the muscle, when lung volume wasincreased by 1.0 liter before stimulation, the rise in Pao amounted to1.75 ± 0.12 cmH₂O. At thehigher volume, Pao/m thereforeaveraged + 0.53 ± 0.05 cmH₂O/g, such that the coefficientof proportionality between the change in triangularis sterni lengthduring passive inflation and Pao/m was the same as that previously obtained for the parasternalintercostal and neck inspiratory muscles. These observations,therefore, confirm that there is a unique relationship between thefractional changes in length of the respiratory muscles, bothinspiratory and expiratory, during passive inflation and theirPao/m. Consequently, the maximal effect of a particular muscle on the lung can be predicted on the basisof its change in length during passive inflation and its mass. Ageometric analysis of the rib cage also established that thelengthening of the canine triangularis sterni during passive inflationis much greater than the shortening of the parasternal intercostalsbecause, in dogs, the costal cartilages slope downward from thesternum.

     

Characterization of expiratory intercostal activity to triangularis sterni in cats

Our purpose was to characterize activity of the intercostal nerve branch innervating the triangularis sterni muscle and the motoneuronal activities comprising this nerve discharge. In decerebrate, vagotomized, paralyzed, and ventilated cats, phasic triangularis sterni neural activity was evident in normocapnia. In most cats, activity did not commence until midexpiration. Activity then rose progressively to terminate at end expiration. Peak neural activities increased in parallel with phrenic activity in hypercapnia and fell in hypocapnia. The progressive increase in triangularis sterni neural activity within each respiratory cycle resulted from recruitment of motoneuronal activities throughout expiration. Once recruited, many motoneurons had a decrementing or constant discharge frequency. In hypercapnia, motoneuronal discharge frequencies increased, and additional activities were recruited. The number of active motoneurons and their discharge frequencies fell in hypocapnia. A similar pattern of motoneuronal activities and responses to stimuli was observed in cats with intact vagi. Factors are considered that may underlie the recruitment pattern of triangularis sterni motoneuronal activities and the inhibition of these in early expiration.     

Diaphragm, genioglossus, and triangularis sterni responses to poikilocapnic hypoxia

Both isocapnic and poikilocapnic hypoxia may elicit a biphasic respiratory response, during which an initial ventilatory stimulation is followed by a reduction in breathing and diaphragm (DIA) electrical activity. To ascertain whether during adulthood other respiratory muscles have biphasic hypoxic responses similar to the DIA, in nine anesthetized cats electromyograms (EMG) were recorded from the DIA, genioglossus (GG), and triangularis sterni (TS) (n = 7) muscles during poikilocapnic hypoxia. DIA and GG EMG started at 60 +/- 4 and 9 +/- 3 units, respectively, during O2 breathing, increased to a maximum of 100 units during the 10-min hypoxic stimulus, and subsequently declined to 81 +/- 6 and 58 +/- 12 units, respectively, by the end of 10 min of hypoxia. The time course of the increase and subsequent decline was similar for the DIA and GG and for GG activity during both inspiration and expiration. Furthermore the degree to which GG EMG declined after reaching its peak activity level correlated with the magnitude of the respective decline in DIA EMG (r = 0.79, P less than 0.02). The TS, in contrast, was maximally active either during O2 breathing or very early during hypoxia, and its activity declined progressively thereafter (to 13 +/- 6% of its peak value at the end of 10 min of hypoxia). The degree to which TS EMG declined did not correlate with the degree to which DIA or GG EMG declined.(ABSTRACT TRUNCATED AT 250 WORDS)     

Ex vivo imaging of motor axon dynamics in murine triangularis sterni explants

We provide a protocol that describes an explant system that allows the dynamics of motor axons to be imaged. This method is based on nerve-muscle explants prepared from the triangularis sterni muscle of mice, a thin muscle that covers the inside of the thorax. These explants, which can be maintained alive for several hours, contain long stretches of peripheral motor axons including their terminal arborizations and neuromuscular junctions. Explants can be prepared from transgenic mouse lines that express fluorescent proteins in neurons or glial cells, which enables direct visualization of their cellular and subcellular morphology by fluorescence microscopy. Time-lapse imaging then provides a convenient and reliable approach to follow the dynamic behavior of motor axons, their surrounding glial cells and their intracellular organelles with high temporal and spatial resolution. Triangularis sterni explants can be prepared in 15 min, imaged ex vivo for several hours and processed for immunohistochemistry in about 2 h.     

Size-related differences in the branching pattern of the motor nerve terminals in triangularis sterni muscle of the mouse

A light microscopy morphometric study was performed in singly innervated synaptic areas of the triangularis sterni muscle of the normal adult Swiss mouse. Investigating mechanisms of the motor nerve growth control, we tested the hypothesis that significant differences in the nerve terminal branching pattern can be detected between different populations of nerve endings classified according to their arborization complexity or size. The main observations of this morphometric study are first, that the mean segment length of the terminal arborization between branch points behaves as an independent variable from the remaining parameters; the mean value of this parameter did not change in nerve endings of differing size and complexity. Secondly, the increase in size of the nerve endings is accompanied by a significant reduction in the mean length of the distal free-end segments. Results are discussed in the context of the possible regulatory mechanisms governing nerve terminal growth and remodelling.     

Role of costal and crural diaphragm and parasternal intercostals during coughing in cats

The inspiratory phase of coughs often consists of large inspired volumes and increased motor discharge to the costal diaphragm. Furthermore, diaphragm electrical activity may persist into the early expiratory portion of coughs. To examine the role of other inspiratory muscles during coughing, electromyograms (EMG) recorded from the crural diaphragm (Dcr) and parasternal intercostal (PSIC) muscles were compared to EMG of the costal diaphragm (Dco) in anesthetized cats. Tracheal or laryngeal stimulation typically produced a series of coughs, with variable increases in peak inspiratory EMGs of all three muscles. On average, peak inspiratory EMG of Dco increased to 346 +/- 60% of control (P less than 0.001), Dcr to 514 +/- 82% of control (P less than 0.0002), and PSIC to 574 +/- 61% of control (P less than 0.0005). Augmentations of Dcr and PSIC EMG were both significantly greater than of Dco EMG (P less than 0.05 and P less than 0.002, respectively). In most animals, EMG of Dco correlated significantly with EMG of Dcr and of PSIC during different size coughs. Electrical activity of all three muscles persisted into the expiratory portions of many (but not all) coughs. The duration of expiratory activity lasted on average 0.17 +/- 0.03 s for Dco, 0.25 +/- 0.06 s for Dcr, and 0.31 +/- 0.09 s for PSIC. These results suggest that multiple respiratory muscles are recruited during inspiration of coughs, and that the persistence of electrical activity into expiration of coughs is not unique to the costal diaphragm.     

Regional intercostal activity during coughing and vomiting in decerebrate cats.

Regional variations in the discharge patterns of the internal and external intercostal muscles of the middle and caudad thorax were studied in decerebrate, spontaneously breathing cats during coughing and vomiting. Coughing, induced by electrical stimulation of the superior laryngeal nerves, consisted of increased and prolonged diaphragmatic activity followed by a burst of abdominal activity. Mid-thoracic external and internal intercostal muscles discharged synchronously with the diaphragm and abdominal muscles, respectively. Caudal external and internal intercostal muscles, however, discharged synchronously with the abdominal muscles. Vomiting, induced by stimulation of the lower thoracic vagi, consisted of a series of synchronous bursts of diaphragmatic and abdominal activity (retching) followed by a prolonged abdominal discharge after the cessation of diaphragmatic activity (expulsion). Caudal external and internal intercostals discharged in phase with diaphragmatic and abdominal activity but both mid-thoracic intercostal muscles discharged out of phase with these muscles. These results indicate major differences in the control and functional roles of intercostal muscles at different thoracic levels during these behaviours.     

Discharge patterns of phrenic motoneurons during fictive coughing and vomiting in decerebrate cats.

In decerebrate, paralyzed, and ventilated cats, we recorded the activity of 100 spontaneously active phrenic motor axons during the increased phrenic discharges characteristic of fictive vomiting (FV) and coughing (FC). During control respiratory cycles, approximately one-half the neurons were recruited in the first decile of inspiration; recruitment continued throughout inspiration. During FV, the duration of phrenic discharge was halved; 20 of 26 motoneurons studied were recruited in the first decile of the burst. During FC, recruitment times did not change compared with control, although the duration of the phrenic burst doubled. Discharge frequencies increased and recruitment order of phrenic motoneurons was virtually unaffected during FC and FV. Limited recruitment of previously inactive neurons in the filaments from which we recorded was found during FV and FC. During FV, 1 previously inactive motoneuron was recruited in 16 filaments containing 25 spontaneously active motor axons. During FC, 3 new motoneurons were recruited in addition to the 64 already active in 35 filaments. Recruitment during FV and FC was absent even when recording from filaments known, on the basis of antidromic activation, to contain inactive motor axons. During FV, 10 of 26 motoneurons began their discharges with doublets (interspike interval < 10 ms); doublets occurred in only 4 of 67 motoneurons during FC. Already active phrenic motoneurons contributed to the intense phrenic activity associated with both respiratory (coughing) and nonrespiratory (vomiting) behavior by increases in discharge frequency, earlier recruitment, and doublets; the contribution of previously quiescent motoneurons remains uncertain.     

The ontogenesis of the manubrium sterni in Tupaia belangeri

The morphogenesis of the manubrium sterni was studied in a series of dated embryos of Tupaia belangeri. In addition to the sternal bands, the "paired suprasternal Anlage" takes part in the shaping of the manubrium sterni as reported by Klima (1968) for other mammals. It forms skeletal elements that mediate between the clavicle and the manubrium: the sternocalvicular ligament and the paired prominence on the dorsal surface of the manubrium, which underlies the clavicles. The paired prominence corresponds to the praeclavium present in some therians. Very probably, the discus articularis of the sternoclavicular articulation of some primates can be attributed to the suprasternal Anlage. There was, however, no indication that the ossa suprasternalia of primates develop from the suprasternal Anlagen: In Tupaia these Anlagen do not form the cranial part of the manubrium. Klima's "unpaired Anlage" develops differently in Tupaia than in other therians. It consists of connective tissue and is not integrated into the manubrium. It presents an insertion surface for the M. pectoralis major, which shifts its origin onto the manubrium, after the sternal bands have fused. The homology of the "unpaired Anlage" and the "pars chondralis interclaviculae" is doubtful.     

Life history and population dynamics of Atriplex triangularis

The dynamics of a population of the stoloniferous Potentilla anserina was studied during four years. The mortality of established ramets followed a log-linear pattern superimposed on a seasonal fluctuation with an increased mortality during the period of vegetative spread. The major external mortality factors were unfavourable weather conditions such as drought and freezing. The turnover rate of the population was slow (31.6 yr). Vegetative spread dominated the recruitment to the population. Seedlings occurred but were exposed to a much higher mortality than vegetative daughter ramets. The juvenile period of seedlings is at least 4 yr in the field. This contrast to vegetative daughter ramets which are potentially flowering and stoloniferous the year after they are established, even if age affects both reproduction and vegetative spread. The density of most of the population is below the level where intraspecific regulative effects operate. However, in very dense stands both stolon and flower production were negatively density dependent.Nomenclature follows Lid (1974), Norsk og Svensk Flora, 2nd ed. Oslo.I am indebted to Lenn Jerling, Peter Torstensson, Anders Telenius and Gudrún Jónsdóttir for their help, and inspiring criticism throughout this study.     

三角叶滨藜的组织培养和植株再生

1　植物名称　三角叶滨藜 (Atriplextriangu laris)。2　材料类别　无菌苗的子叶。3　培养条件　培养基 :( 1 )MS + 6 BA 1mg·L- 1(单位下同 ) + 2 ,4 D 1 ;( 2 )MS + 6 BA 1 +NAA 1 ;( 3)MS + 6 BA 1 +IBA 1 ;( 4 )MS + 6 BA 1 +IAA1 ;( 5 )MS + 6 BA 0 .5 +IAA 0 .1 +GA 0 .2 ;( 6)MS + 6 BA 0 .5 +IAA 0 .4+GA 2 ;( 7)MS + 6 BA1 +IAA 0 .2 +GA 2 ;( 8)MS + 6 BA 1 +IAA 0 .4+GA 2 ;( 9) 1 /2MS +IAA 2 ;( 1 0 ) 1 /2MS。上述培养基 ( 1 )～ ( 4 )加CH(水解酪蛋白 ) 2 0 0、45 g·L- 1 蔗糖 ,( 5 )～ ( 8…     

Regeneration of Oxalis triangularis ssp. triangularis from suspension cells cultured in three different systems (solid, liquid-flask and bioreactor cultures)

In attemps to establish in vitro cultures of Oxalis triangularis ssp. Triangularis, the explants of leaves, petioles, bulb scales and suspension cells derived from regenerated bulbs were examined using solid (petri dish), liquid-flask and bioreactor cultures. Only bulb-derived suspension cells were able to regenerate in all culture systems. The liquid-flask and bioreactor cultures supported organogenesis and yielded larger amount of buds than solid culture. The inclusion of AC in the culture medium delayed bud initiation but promoted plantlet development by minimising callusing of the buds. Morphological differences in regenerated plantlets affected by AC, such as the length and diameter of the petiole, leaf unfolding and the development of a red colour development on leaves and petioles, varied with the culture systems. Upon transfer to pots normal plants were recovered from buds regenerated in various culture systems. Received: 18 August 1998 / Revision received: 27 October 1998 / Accepted: 20 November 1998     

Neuronal differentiation in the thalamic area triangularis of a lizard

Development of neurons in the area triangularis of Gallotia galloti was investigated in Golgi-impregnated brain tissue. Four major neuronal types present in adults were found to originate from two migratory neuroblast types, which were followed from embryonic stage S.32. One type has a thick main medial process, whereas the second type has a long main lateral process. As they migrate toward the periphery of the nucleus, morphological characteristics of maturation appear, including growth cones, filopodia, and outgrowth of axons. Neuroblasts with a main lateral process differentiate into two immature neuronal types, bipolars and pyramidals, observed at S.33 and thereafter. The neuroblasts with a main medial process undergo some somatic translocation through a transitory tangential shaft. Then they develop into monopolar immature forms with a long varicose medial, process, appearing from S.36. onward. Immature bipolar neurons do not experience great changes in their dendritic arborization during development to the adult stage, but pyramidals and monopolars undergo a rapid development of the dendritic tree after S.36. By S.38 archetypes of adult neuronal forms are established. Hairlike appendages first appear on neurons at S.36 They decrease suddenly in S.38 and then proliferate in S.39 when spines first appear. Around the time of hatching, the hairlike appendages begin to disappear and spines become established. Reduction of spines occurs after hatching and continues to the adult stage. Possible influences of several external factors on neuronal maturation are discussed.     

Triangularis sterni: a primary muscle of breathing in the dog

The isolated action, pattern of neural activation, and mechanical contribution to eupnea of the triangularis sterni (transversus thoracis) muscle were studied in supine anesthetized dogs. Linear displacement transducers were used to measure the axial displacements of the ribs and sternum. Tetanic stimulation of the triangularis sterni in the apneic animal caused a marked caudal displacement of the ribs, a moderate cranial displacement of the sternum, and a decrease in lung volume. During quiet breathing, there was invariably a rhythmic activation of the muscle in phase with expiration that was independent of the presence or absence of activity in the abdominal and internal interosseous intercostal muscles. This phasic expiratory activity in the triangularis sterni was of large amplitude and caused the ribs to be more caudal and the sternum to be more cranial during the spontaneous expiratory pause than during relaxation. Additional studies on awake animals showed that rhythmic activation of the triangularis sterni occurs in all body positions and is not caused by anesthesia. These findings indicate that expiration in the dog is not a passive process and that the end-expiratory volume of the rib cage is not determined by an equilibrium of static forces alone. Rather, it is actively determined and maintained below its relaxation volume by contraction of the triangularis sterni throughout expiration. The use of this muscle is likely to facilitate inspiration by increasing the length of the parasternal intercostals and taking on a portion of their work.