Tracheal length changes during zebra finch song and their possible role in upper vocal tract filtering

Sounds produced in the avian vocal organ may be modified by filter properties of the upper vocal tract. Possible mechanisms to actively control filter characteristics include movements of the beak, tongue, and larynx and adjustments of tracheal length. We investigated whether length changes of the trachea are a likely mechanism for adjusting upper vocal tract filter properties during song in the zebra finch (Taeniopygia guttata). Tracheal length was monitored at the basal end using sonomicrometry and was recorded together with subsyringeal air sac pressure and acoustic output. Tracheal shortening occurred at the onset of song bouts, and during each motif the tracheal length decreased during expiratory pressure pulses and increased during the short inspirations. A bilateral tracheal syringeal nerve cut confirmed that the initial shortening at the onset of the song bout is an active shortening of the trachea (i.e., mediated by syringeal muscle activity). The modulation of length during the motif was not affected by the denervation and is most likely driven by the pressurization of the interclavicular air sac. The absolute length change during the motif was small (<0.2 mm) and not clearly related to acoustic features of the song. For example, some high-frequency syllables, which are generated during inspiration, were accompanied by tracheal elongation. Because this elongation shifts tube resonances to lower frequencies, it is inconsistent with an active adjustment of length to enhance high frequency sounds. The small magnitude and inconsistent nature of dynamic tracheal length changes during song make it unlikely that they significantly affect vocal tract filter properties if the trachea is modeled as a rigid tube.     

Migrate small,sound big: functional constraints on body size promote tracheal elongation in cranes

Organismal traits often represent the outcome of opposing selection pressures. Although social or sexual selection can cause the evolution of traits that constrain function or survival (e.g. ornamental feathers), it is unclear how the strength and direction of selection respond to ecological shifts that increase the severity of the constraint. For example, reduced body size might evolve by natural selection to enhance flight performance in migratory birds, but social or sexual selection favouring large body size may provide a countervailing force. Tracheal elongation is a potential outcome of these opposing pressures because it allows birds to convey an auditory signal of exaggerated body size. We predicted that the evolution of migration in cranes has coincided with a reduction in body size and a concomitant intensification of social or sexual selection for apparent large body size via tracheal elongation. We used a phylogenetic comparative approach to examine the relationships among migration distance, body mass and trachea length in cranes. As predicted, we found that migration distance correlated negatively with body size and positively with proportional trachea length. This result was consistent with our hypothesis that evolutionary reductions in body size led to intensified selection for trachea length. The most likely ultimate causes of intensified positive selection on trachea length are the direct benefits of conveying a large body size in intraspecific contests for mates and territories. We conclude that the strength of social or sexual selection on crane body size is linked to the degree of functional constraint.     

Pulmonary lesions in disseminated visceral coccidiosis of sandhill and whooping cranes

Fifty cranes, consisting of 46 sandhill (Grus canadensis) and four whooping cranes (Grus americana), were studied. Eighteen sandhill cranes and the four whooping cranes were naturally infected with disseminated visceral coccidiosis (DVC). The remaining sandhill cranes were chicks experimentally infected with oocysts of Eimeria reichenowi and/or E. gruis; five chicks served as controls. There were no clinical signs attributed to respiratory infection. Necropsy of naturally infected adult birds revealed nodules in many organs, including the lung, air sacs, trachea and nares. Artificially infected sandhill cranes and the whooping crane chicks that died from DVC had congestion and consolidated areas in the lung with frothy fluid in the airways. Grossly visible nodules were observed from 10 days postinoculation. Granulomatous pneumonia and tracheitis were observed with light microscopy. Lesions were associated with merogonic and gametogonic stages of eimerian coccidia. Granulomas and granulomatous foci contained parasitized large mononuclear cells. Merogonic stages were seen in lymphoid cells by ultrastructural examination. Oocysts were observed in the trachea and bronchial mucosa and admixed with exudate in the airways, indicating that crane eimerians can complete their life cycle at these sites. Of the few eimeriid coccidia that have extraintestinal stages of development in birds and mammals, only the species in cranes complete their life cycle in both the digestive and respiratory tracts.     

In vivo loads on airway smooth muscle: the role of noncontractile airway structures.

The degree of airway smooth muscle contraction and shortening that occurs in vivo is modified by many factors, including those that influence the degree of muscle activation, the resting muscle length, and the loads against which the muscle contracts. Canine trachealis muscle will shorten up to 70% of starting length from optimal length in vitro but will only shorten by around 30% in vivo. This limitation of shortening may be a result of the muscle shortening against an elastic load such as could be applied by tracheal cartilage. Limitation of airway smooth muscle shortening in smaller airways may be the result of contraction against an elastic load, such as could be applied by lung parenchymal recoil. Measurement of the elastic loads applied by the tracheal cartilage to the trachealis muscle and by lung parenchymal recoil to smooth muscle of smaller airways were performed in canine preparations. In both experiments the calculated elastic loads applied by the cartilage and the parenchymal recoil explained in part the limitation of maximal active shortening and airway narrowing observed. We conclude that the elastic loads provided by surrounding structures are important in determining the degree of airway smooth muscle shortening and the resultant airway narrowing.     

Novel mechanisms of tube-size regulation revealed by the Drosophila trachea

The size of various tubes within tubular organs such as the lung, vascular system and kidney must be finely tuned for the optimal delivery of gases, nutrients, waste and cells within the entire organism. Aberrant tube sizes lead to devastating human illnesses, such as polycystic kidney disease, fibrocystic breast disease, pancreatic cystic neoplasm and thyroid nodules. However, the underlying mechanisms that are responsible for tube-size regulation have yet to be fully understood. Therefore, no effective treatments are available for disorders caused by tube-size defects. Recently, the Drosophila tracheal system has emerged as an excellent in vivo model to explore the fundamental mechanisms of tube-size regulation. Here, we discuss the role of the apical luminal matrix, cell polarity and signaling pathways in regulating tube size in Drosophila trachea. Previous studies of the Drosophila tracheal system have provided general insights into epithelial tube morphogenesis. Mechanisms that regulate tube size in Drosophila trachea could be well conserved in mammalian tubular organs. This knowledge should greatly aid our understanding of tubular organogenesis in vertebrates and potentially lead to new avenues for the treatment of human disease caused by tube-size defects.     

Serrano (Sano) Functions with the Planar Cell Polarity Genes to Control Tracheal Tube Length

Epithelial tubes are the functional units of many organs, and proper tube geometry is crucial for organ function. Here, we characterize serrano (sano), a novel cytoplasmic protein that is apically enriched in several tube-forming epithelia in Drosophila, including the tracheal system. Loss of sano results in elongated tracheae, whereas Sano overexpression causes shortened tracheae with reduced apical boundaries. Sano overexpression during larval and pupal stages causes planar cell polarity (PCP) defects in several adult tissues. In Sano-overexpressing pupal wing cells, core PCP proteins are mislocalized and prehairs are misoriented; sano loss or overexpression in the eye disrupts ommatidial polarity and rotation. Importantly, Sano binds the PCP regulator Dishevelled (Dsh), and loss or ectopic expression of many known PCP proteins in the trachea gives rise to similar defects observed with loss or gain of sano, revealing a previously unrecognized role for PCP pathway components in tube size control.     

四种鹤的胸骨和肩带骨比较

本文对国家一类保护动物丹顶鹤、白枕鹤、白头鹤、白鹤的胸骨与肩带骨进行了比较,阐明白鹤与鹤属的3种鹤在骨胳特征上的重要差别,进一步论证把白鹤从鹤属分离出来与肉垂鹤合为一个属是正确的。本文首次就胸骨和肩带骨特征提出4种鹤的检索表。文中附有胸骨和肩带骨的量度和12幅鹤的胸骨图。     

Uncinate Process Length in Birds Scales with Resting Metabolic Rate

A fundamental function of the respiratory system is the supply of oxygen to meet metabolic demand. Morphological constraints on the supply of oxygen, such as the structure of the lung, have previously been studied in birds. Recent research has shown that uncinate processes (UP) are important respiratory structures in birds, facilitating inspiratory and expiratory movements of the ribs and sternum. Uncinate process length (UPL) is important for determining the mechanical advantage for these respiratory movements. Here we report on the relationship between UPL, body size, metabolic demand and locomotor specialisation in birds. UPL was found to scale isometrically with body mass. Process length is greatest in specialist diving birds, shortest in walking birds and intermediate length in all others relative to body size. Examination of the interaction between the length of the UP and metabolic demand indicated that, relative to body size, species with high metabolic rates have corresponding elongated UP. We propose that elongated UP confer an advantage on the supply of oxygen, perhaps by improving the mechanical advantage and reducing the energetic cost of movements of the ribs and sternum.     

Perception of Vocal Tract Resonances by Whooping Cranes Grus americana

Although formants (vocal tract resonances) can often be observed in avian vocalizations, and several bird species have been shown to perceive formants in human speech sounds, no studies have examined formant perception in birds' own species-specific calls. We used playbacks of computer-synthesized crane calls in a modified habituation—dishabituation paradigm to test for formant perception in whooping cranes ( Grus americana ). After habituating birds to recordings of natural contact calls, we played a synthesized replica of one of the habituating stimuli as a control to ensure that the synthesizer worked adequately; birds dishabituated in only one of 13 cases. Then, we played the same call with its formant frequencies shifted. The birds dishabituated to the formant-shifted calls in 10 out of 12 playbacks. These data suggest that cranes perceive and attend to changes in formant frequencies in their own species-specific vocalizations, and are consistent with the hypothesis that formants can provide acoustic cues to individuality and body size.     

Constructing an organ: the Drosophila salivary gland as a model for tube formation

Tubes are required in metazoans to transport the liquids and gases that sustain life. The conservation of molecules and mechanisms involved in tube formation suggests that what we learn by studying simple systems will apply to related processes in higher animals. Studies over the past 10 years have revealed the molecules that specify cell fate in Drosophila salivary gland and the cellular events that mediate tube morphogenesis. Here, we discuss how anterior-posterior and dorsal-ventral patterning information specifies both the position of salivary-gland primordia and how many cells they contain. We examine the transformation of a polarized epithelial sheet into an elongated, unbranched tube, and the intrinsic and extrinsic factors that influence the final position of the salivary gland.     

Distal projection of insufflated gas during tracheal gas insufflation.

Tracheal gas insufflation (TGI) flushes expired gas from the ventilator circuitry and central airways, augmenting CO2 clearance. Whereas a significant portion of this washout effect may occur distal to the injection orifice, the penetration and mixing behavior of TGI gas has not been studied experimentally. We examined the behavior of 100% oxygen TGI injected at set flow rates of 1-20 l/min into a simulated trachea consisting of a smooth-walled, 14-mm-diameter tube. Models incorporating a separate coaxial TGI injector, a rough-walled trachea, and a bifurcated trachea were also studied. One-hundred percent nitrogen, representing expiratory flow, passed in the direction opposite to TGI at set flow rates of 1-25 l/min. Oxygen concentration within the "trachea" was mapped as a function of axial and radial position. Three consistent findings were observed: 1) mixing of expiratory and TGI gases occurred close to the TGI orifice; 2) the oxygenated domain extended several centimeters beyond the endotracheal tube, even at high-expiratory flows, but had a defined distal limit; and 3) more distally from the site of gas injection, the TGI gas tended to propagate along the tracheal wall, rather than as a central projection. We conclude that forward-directed TGI penetrates a substantial distance into the central airways, extending the compartment susceptible to CO2 washout.     

Functional anatomy of the vagal innervation of the cervical trachea of the dog.

The canine cervical trachea has been used for numerous studies regarding the neural control of tracheal smooth muscle. The purpose of the present study was to determine whether there is lateral dominance by either the left or right vagal innervation of the canine cervical trachea. In anesthetized dogs, pressure in the cuff of the endotracheal tube was used as an index of smooth muscle tone in the trachea. After establishment of tracheal tone, as indicated by increased cuff pressure, either the right or left vagus nerve was sectioned followed by section of the contralateral vagus. Sectioning the right vagus first resulted in total loss of tone in the cervical trachea, whereas sectioning the left vagus first produced either a partial or no decrease in tracheal tone. After bilateral section of the vagi, cuff pressure was recorded during electrical stimulation of the rostral end of the right or left vagus. At the maximum current strength used, stimulation of the left vagus produced tracheal constriction that averaged 28.5% of the response to stimulation of the right vagus (9.0 +/- 1.8 and 31.6 +/- 2.5 mmHg, respectively). In conclusion, the musculature of cervical trachea in the dog appears to be predominantly controlled by vagal efferents in the right vagus nerve.     

Tracheal compliance and limit flow rate changes in a murine model of asthma

Trachea is the unique passage for air to flow in and out. Its tone is of importance for the respiration system. However, investigation on how tracheal tone changes due to asthma is limited. Aiming at studying how the mechanical property changes due to asthma as well as the compliance and flow limitation, the following methods are adopted. Static and passive pressure-volume tests of rats' trachea of the asthmatic and control groups are carried out and a new type of tube law is formulated to fit the experimental data, based on which changes of compliance and limit flow rate are investigated. In order to give explanation to such changes, histological examinations with tracheal soft tissues are made. The results show that compliance, limit flow rate and material constants included in the tube law largely depend on the longitudinal stretching ratio. Compared with the control group, the tracheal compliance of asthmatic animals decreases significantly, which results in an increased limit flow rate. Histological studies indicate that asthma can lead to hyperplasia/hypertrophy of smooth muscle cells, and increase elastin and collagen fibres in the muscular membrane. Though decreasing compliance increases sta- bility, during the onset of asthma, limit flow rate is much smaller due to the lower transmural pressure. Asthma leads to a stiffer trachea and the obtained results reveal some aspects relevant to asthma-induced tracheal remodelling.     

Quantitative measurement of smooth muscle shortening in isolated pig trachea

Matched porcine tracheal rings were exposed to theophylline and increasing doses of carbachol in Krebs solution. Histological sections of each ring were traced and each of the following dimensions measured: the external perimeter (Pe) and external area (Ae) defined by the outer border of smooth muscle and inner surface of cartilage, and the internal perimeter (Pi) and internal area (Ai) defined by the luminal surface of the epithelium and the muscle length (L) along its outer border. Absolute wall area (WA = Ae - Ai) and relative wall area (PW = WA/Ae) were calculated. Carbachol-treated tracheal ring dimensions were compared with those of their matched theophylline-treated rings. In tracheal rings with intact cartilage, maximal smooth muscle shortening of 44% was achieved with 10(-2) M carbachol. In tracheal rings in which anterior and posterior segments of cartilage were excised, the trachealis muscle passively shortened by 20% and maximal shortening (10(-3) M carbachol) was 57%. Although Ai decreased with maximal smooth muscle shortening, there were no changes in the length of Pi or in WA. These data show that the cartilage in the porcine trachea exerts both a preload that passively stretches the trachealis muscle and an afterload that limits maximal smooth muscle shortening.     

Ventilatory failure induced by tracheal banding in the hamster.

Previous animal models of hypercapnic ventilatory failure are limited in that the resistive load has only been applied acutely and often in anesthetized animals. We therefore developed a chronic animal model of hypercapnic ventilatory failure by increasing airway resistance via tracheal banding over several days. To test the efficacy of this model, we compared arterial blood gases, pulmonary function, and internal area of the trachea 6 days after the banding or sham procedure in 20 hamsters. Six days later, banded animals had an increased airway resistance as indicated by a 66% reduction in internal cross-sectional area of the trachea and a 6.5-fold increase in pulmonary resistance compared with control hamsters. The increased airway resistance resulted in a severe respiratory acidosis and hypoxemia in the awake banded hamsters. Banded hamsters were also hyperinflated. This animal model will be useful for investigating the various mechanisms that contribute to hypercapnic ventilatory failure and interventions that may promote recovery.     

Tracheal dimensions at full inflation and deflation in adolescent twins

Tracheal dimensions at total lung capacity (TLC) and residual volume (RV) were analyzed roentgenographically in 17 pairs of male adolescent twins (mean age 16.3 yr; 12 monozygotic pairs and 5 dizygotic pairs). Genetic factors dominated environmental traits in intra- as well as extrathoracic tracheal width at RV. Extrathoracic tracheal width at TLC was also governed by genetic components. Intrathoracic tracheal depth (anteroposterior diameter), length, and cross-sectional area did not seem to be genetically controlled at TLC and RV. Intrathoracic tracheal cross-sectional area increased by 14.4% and became more elliptical from RV to TLC, owing mainly to an increase in tracheal depth (16.7%). Increments from RV to TLC in tracheal depth but not width correlated with increases in lung width, depth, and height. Intrathoracic trachea was elongated 14% in association with increase in lung height from RV to TLC. At TLC, extrathoracic tracheal width was larger than intrathoracic tracheal width, but this dimension did not differ at RV. These results indicate that genetic factors influence, at least at RV, the tracheal rings more strongly than membranous parts. Intrathoracic tracheal depth but not width increases during inspiration in accordance with increase in lung volume. Extrathoracic tracheal width widens more than intrathoracic trachea from RV to TLC.     

Effect of positional changes of anatomic structures on upper airway dilating muscle shortening during electro- and chemostimulation.

Positional changes of anatomic structures surrounding the upper airway are known to affect pharyngeal mechanics and collapsibility. We hypothesized that these alterations also affect the ability of the upper airway dilator muscles to enlarge the pharynx by altering their ability to shorten when activated. Using sonomicrometry, we evaluated in seven anesthetized dogs the effects of changes in tracheal and head position on the length of the genioglossus (GG) and the geniohyoid (GH) and the effects of these positional changes on the magnitude of shortening of the two muscles in response to electro- (ES) and chemostimulation (CS). Caudal traction of the trachea lengthened the GG and GH in all dogs, whereas cranial displacement of the trachea and flexion of the head to a vertical position shortened the muscles. Compared with the magnitude of ES-induced shortening in the neutral position, ES-induced shortening of the GG was 144.7 +/- 14.6, 49.3 +/- 4.3, and 33.5 +/- 11.6% during caudal and cranial displacement of the trachea and during head flexion, respectively. Similar effects of the positional changes were found for the GH, as well as for both muscles during respiratory stimulation with P(CO2) of 90 Torr at the end of CO(2) rebreathing, although inspiratory muscle shortening during CS reached only one-quarter to one-third of the magnitude observed during ES. We conclude that positional alterations of anatomic structures in the neck have a dramatic effect on the magnitude of shortening of the activated GG and GH, which may reduce substantially their ability to protect pharyngeal patency.     

Ultrastructure of the mouse tracheal epithelium

The ultrastructure of mouse tracheal epithelium was examined. The three cell types, basal cells, ciliated cells and goblet cells, described for other mammalian trachea were found to be present although goblet cells occurred only rarely. A cell type, termed the nonciliated cell, not described in other mammalian trachea was frequently found in mouse tracheal epithelium. These cells contained abundant smooth and rough endoplasmic reticulum, free ribosomes, a large Golgi complex, and many mitochondria. There were many vesciles containing an electron dense material near the luminal surface of these cells; these cells were positive for PAS. These features suggested a secretory function for the cells. This, along with the scarcity of goblet cells, suggested that the nonciliated cells of mouse tracheal epithelium fulfill the function of the goblet cells found in other mammalian trachea.     

Zum Zwischenstopp nach Ost‐Frankreich

Stopover in France The Lac du Der and És?tang de Lindre belong to a group of lakes located in France. Lots of migratory birds especially cranes choose this region for a stopover every year. Not only because of the fascinating spectacle when the cranes land at the Lac du Der at dusk, but also because of the many raptors and aquatic birds living in this region it is worth visiting the lakes for experts aswell as for laypersons.