Zur Morphogenese des respiratorischen Epithels der Tracheenkiemen bei Larven der Limnephilini KOL. (Insecta,Trichoptera)

Zusammenfassung Das respiratorische Epithel der Tracheenkiemen ist durch ein hochgeordnetes Tracheolengerüst charakterisiert. Die Tracheolen liegen parallel zur Längsachse der fadenförmigen Tracheenkiemen dicht unter der Cuticula in statistisch gleichmäßigem Abstand zueinander. Die Regelmäßigkeit dieses subcuticularen Tracheolengerüsts weist auf ein physiologisch optimal arbeitendes System hin. Der Abstand zwischen zwei Tracheolen ist sehr wahrscheinlich gleich dem doppelten Radius der tracheolaren Einzugsgebiete. Auf diese Weise wird bei einem Minimum an Tracheolenmaterial der gesamte diffundierende Sauerstoff der respiratorischen Oberfläche von den Tracheolen erfaßt. Die Morphogenese dieser Strukturregelmäßigkeit wird während der larvalen Entwicklung verfolgt. Dabei zeigt sich, daß mit jeder Häutung zahlreiche neue Tracheolen in das respiratorische Epithel geordnet eingebaut werden und die Abstände zwischen den Tracheolen in Korrelation zum Radius der Tracheolen von Larvenstadium zu Larvenstadium geregelt abnehmen.

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Morphogenesis of the respiratory epithelium in the tracheal gills of larval Limnephilini KOL. (Insecta, Trichoptera)

Summary The respiratory epithelium of the tracheal gills of the larval Limnephilini KOL. (Insecta, Trichoptera) is characterized by a highly organized tracheolar framework. The tracheoles are found parallel to the longitudinal axis of the thread-like tracheal gills and lie closely underneath the cuticle at statistically uniform distances. The regular distribution of the subcuticular tracheoles represents an optimum physiological system with the tracheolar interspace probably corresponding to twice the radius of the tracheolar catchment area. This arrangement ensures that all oxygen diffusing across the respiratory gill surface is taken up by the tracheoles with a minimum of tracheolar material. The morphogenesis of this regular distribution was studied during the larval development. With each moult numerous new tracheoles are added to the regular distribution. The distances between the tracheoles decrease regularly in correlation to the decreasing radius of the tracheoles from one larval stage to the next.

     

Ultrastructural observation on the rectal gills of Zygoptera larva.

The zygoptera larvae have tracheal gills. Both outer and inner epicuticles of damselfly larvae (Zygoptera) are pierced by pores of about 200...250 A thickness. The pores are distributed at a distance of about 500...600 A apart. Their endings are noticed upto the lower portion of exocuticle. The endocuticle is a big area in which the respiratory epithelium have flat apical and basal sides, plasma membrane is not folded. Glycogen granules are distributed in the cytoplasm and a few mitochondria are noticed. The tracheal brances and the cytoplasm of the respiratory epithelium come in contact with the epicuticle. The distribution of tracheoles, with surplus of tracheoles in a surface outside the hypodermis are close to the sub-cuticular area is found. The gills of Zygoptera appear to be merely respiratory organs having respiratory epithelium and the ions and water are resorbed from the modified rectal epithelium.     

Tracheation of abdominal ganglia and cerci in the house cricket Acheta domesticus (Orthoptera,Gryllidae)

Patterns of tracheation in the abdominal central nervous system and the cerci of Acheta domesticus are described from whole mounts, and light and electron microscopy. The tracheal supply of the ganglia is derived from ventral longitudinal tracheal trunks which have segmental connections to the spiracels. Each abdominal ganglion is served by a single pair of tracheal trunks, except the terminal ganglion, which has two pairs. Within the ganglia, tracheoles occur principally in association with glia-rich areas of the neuropile. We suggest that the respiratory exchange may be concentrated in the cell bodies of neurons and glia. Each cercus has a tracheal supply in paralle with a large air sac which, it is suggested, serves to lighten the cercus, functions as a resonator for sound reception, or facilitates tidal flow of hemolymph and postecdysial expansion of the cercus. No tracheae run continuously between ganglia or between the terminal ganglion and the cerci, and they do not appear to have a potential role as a contact guidance pathway for cercal nerve growth.     

Transition regime diffusion and the structure of the insect tracheolar system

A model is developed for the diffusion of oxygen along insect tracheoles. It is shown that reduction of the effective tracheolar diameter is advantageous until this quantity approaches the mean free path of diffusing oxygen molecules, and that, once this transition regime has been entered, further reduction of tracheolar diameter is disadvantageous. This finding offers a plausible explanation of Weis-Fogh's observation that the respiratory pathway in insects continues to branch until the smallest ‘twigs’ approach a mean free path in diameter.     

The tracheae of water mites

The water mites of standing waters have evolved a novel respiratory system consisting of numerous independent tracheae of tracheolar dimensions. Each trachea has a portion of its length lying directly under the cuticle and one or both ends of the trachea turn into the body to supply some organ. There is no fusion of tracheae to form trunks. Areas of dense tracheation dorsal to the legs supply the leg muscles, and sometimes there is a distinct area of the venter that supplies the muscles of the mouthparts.     

Reinforced tracheoles in three firefly lanterns: Further reflections on specialized tracheoles

The structures of the lantern tracheoles of three genera of flashing fireflies are compared. All three genera have stiff, reinforced tracheoles which resist folding or collapsing under conditions which flatten more typical tracheoles. This common specialization supports the hypothesis that the tracheoles play a major role in flash control in these fireflies, especially as the morphological basis of the stiffening is different in the three genera. Study of the tracheoles of other tissues reveals that there is great variety in structure and flexibility of these vessels from tissue to tissue and organism to organism, suggesting that tracheolar specialization may be a general phenomenon, with the fine structure of these air tubes being tailored to the particular demands and conditions of the tissues in which they are found.     

Permeable sites in the firefly lantern tracheal system: Use of osmium tetroxide vapor as a tracer

Flashing fireflies were permitted to breathe osmium tetroxide vapor, after which the lanterns were removed and the sites of absorption of the osmium into the tissues were detected in two ways: (1) by sonication to remove soft tissues, that is, those that had not been fixed by the osmium gas, and (2) by intensification with thiocarbohydrazide and silver nitrate, in a modification of the osmium–thiocarbohydrazide–osmium (OTO) stain technique. The results of both procedures indicate that the gas first enters into the tissues at the level of the tracheoles. These findings may be interpreted as underscoring the importance of the tracheolar cell and the tracheal end organ in the control of oxygen entry into the lantern tissues, and the implications of the results in the oxygen regulation theory of flash control are discussed.     

Corrosion casts as a method for investigation of the insect tracheal system

Summary The tracheal systems of five insect species (two species of ants, worker bee, housefly and the cabbage butterfly) have been studied by scanning electron microscopy of corrosion casts. This technique, which is commonly used for the investigation of vertebrate vasculature, is adapted to demonstrate the ultrastructure of the insect respiratory organ. The problem of filling a blind ending system was solved by injecting the resin Mercox into the evacuated tracheae through a thoracal spiracle. After polymerization of the resin, the tissue was digested enzymatically and chemically. The three-dimensional structure of the tracheal system was investigated by scanning electron microscopy. The technique used here displays for the first time the complex morphology of the entire tracheal system in fine detail, especially the structure of spiracles, airsacs, tracheae and tracheoles. Smooth-walled terminal tracheoles show up in flight muscles. The finest tracheoles that could be identified have diameters of approximately 70 nm. This approaches the finest tracheoles portrayed by transmission electron micrographs.     

雷氏黄萤幼虫呼吸系统和呼吸行为

研究雷氏黄萤Luciola leii Fu and Ballantyne幼虫的呼吸系统及其呼吸行为。结果表明:雷氏黄萤幼虫的呼吸系统中只有气管无气囊。前胸、中胸和后胸均分布有气门,无气管鳃,腹部1~8节分布有气门和气管鳃,气门腔基部和气管鳃基部相连,呈"√"状,气管鳃内气管与气门气管相连通。雷氏黄萤幼虫的呼吸行为分为3种:利用胸部气门呼吸、腹部气门呼吸和气管鳃呼吸,其中以腹部气门呼吸为主。     

Tghe tracheation of the wings of adult Lepidoptera

There is much variation in the extent of tracheation of the wings of adult Lepidoptera. This is surveyed. Significantly more tracheal branching is found in species with pupal cocoons. No correlation of tracheation with wing length is apparent. The concentrations of oxygen and carbon dioxide in the air inside cocoons of Antheraea pernyi Guér. (Saturniidae) before emergence are measured. Oxygen supply is probably not limited by the cocoon. Respiration rates per unit mass of wings of A. pernyi are similar to those of the whole, resting adult. The possible adaptive significance of this tracheation and its variation is discussed.     

Microscale Gaseous Slip Flow in the Insect Trachea and Tracheoles

An analytical investigation into compressible gas flow with slight rarefactions through the insect trachea and tracheoles during the closed spiracle phase is undertaken, and a complete set of asymptotic analytical solutions is presented. We first obtain estimates of the Reynolds and Mach numbers at the channel terminal ends where the tracheoles directly deliver respiratory gases to the cells, by comparing the magnitude of the different forces in the compressible gas flow. The 2D Navier–Stokes equations with a slip boundary condition are used to investigate compressibility and rarefied effects in the trachea and tracheoles. Expressions for the velocity components, pressure gradients and net flow inside the trachea are then presented. Numerical simulations of the tracheal compressible flow are performed to validate the analytical results from this study. This work extends previous work of Arkilic et al. (J Microelectromech Syst 6(2):167–178, 1997) on compressible flows through a microchannel. Novel devices for microfluidic compressible flow transport may be invented from results obtained in this study.     

The structure and function of the gill tufts in larval Amphipterygidae (Odonata:zygoptera)

Larvae of the subfamily Amphipteryginae (Odonata) bear a tuft of tracheal gills on either side of the anus. The two tufts are derived from the laminae sub-anales, and are protected by the non-respiratory epiproct and paraprocts, and by plates derived from the cerci, lamina supra-analis or the lamina sub-analis itself. Each is approximately 1 mm long in mature larvae and comprises a series of repeatedly branching filaments, the terminal twigs of which are 5 to 10 μ in diameter. The total surface area of the tufts is approximately 5.0 mm² in mature larvae of Devadatta, and more in the larvae of Pentaphlebia and Rimanella. Each tuft is connected by a large trachea to the longitudinal tracheal trunk. This large trachea divides many times, eventually forming a dense palisade of tracheoles in the epidermis of the filaments, immediately beneath the thin investing cuticle.     

Moulting of insect tracheae captured by light and electron-microscopy in the metathoracic femur of a third instar locust Locusta migratoria

The insect tracheal system is an air-filled branching network of internal tubing that functions to exchange respiratory gases between the tissues and the environment. The light and electron-micrographs presented in this study show tracheae in the process of moulting, captured from the metathoracic hopping femur of a juvenile third instar locust (Locusta migratoria). The images provide evidence for the detachment of the cuticular intima from the tracheal epithelial cells, the presence of moulting fluid between the new and old cuticle layers, and the withdrawal of the shed cuticular lining through larger upstream regions of the tracheal system during moulting. The micrographs also reveal that the cuticular intima of the fine terminal branches of the tracheal system is cast at ecdysis. Therefore, the hypothesis that tracheoles retain their cuticle lining at each moult may not apply to all insect species or developmental stages.     

Association of tracheal placodes with leg primordia in Drosophila and implications for the origin of insect tracheal systems

Adaptation to diverse habitats has prompted the development of distinct organs in different animals to better exploit their living conditions. This is the case for the respiratory organs of arthropods, ranging from tracheae in terrestrial insects to gills in aquatic crustaceans. Although Drosophila tracheal development has been studied extensively, the origin of the tracheal system has been a long-standing mystery. Here, we show that tracheal placodes and leg primordia arise from a common pool of cells in Drosophila, with differences in their fate controlled by the activation state of the wingless signalling pathway. We have also been able to elucidate early events that trigger leg specification and to show that cryptic appendage primordia are associated with the tracheal placodes even in abdominal segments. The association between tracheal and appendage primordia in Drosophila is reminiscent of the association between gills and appendages in crustaceans. This similarity is strengthened by the finding that homologues of tracheal inducer genes are specifically expressed in the gills of crustaceans. We conclude that crustacean gills and insect tracheae share a number of features that raise the possibility of an evolutionary relationship between these structures. We propose an evolutionary scenario that accommodates the available data.     

Morphometric analysis of the larval branchial chamber in the dragonfly Aeshna cyanea Müller (Insecta, Odonata, Anisoptera)

The aquatic larvae of anisopteran dragonflies possess tracheal gills located in the rectum. Using stereological methods, we estimated the morphometric diffusing capacity for oxygen (D(MO2)) across the gill epithelium, i.e., from rectal water to the gill tracheoles, in the larvae of Aeshna cyanea. A 271-mg larva has a total branchial surface area of approximately 12 cm(2). Tracheoles make up 6% of the epithelial volume of the gills; the harmonic mean thickness of the water-tracheolar diffusion barrier is 0.27 microm and consists mainly of cuticle. The calculated D(MO2) is 23.0 microl min(-1) g(-1) kPa(-1), which, using published values for oxygen consumption in a similar species, would result in a mean driving pressure of 0.2 kPa at rest and 1.3 kPa during activity. Since these driving pressures are similar to those reported for other arthropods, we conclude that the D(MO2) of the gill is not rate-limiting for aerobic metabolism in Aeshna cyanea larvae. J Morphol. 261:81-91, 2004.     

Stereological determination of tracheal volume and diffusing capacity of the tracheal walls in the stick insect Carausius morosus (Phasmatodea, Lonchodidae)

First instars of Carausius morosus provide a good model for morphometric evaluation of the diffusing capacity between the tracheal system and hemolymph: air sacs are lacking, tracheoles do not penetrate the organs and muscles, and entire animals can be evaluated electron microscopically without subsampling. The tracheal volume makes up 1.3% of the volume of the whole insect excluding appendages. We calculated the lateral diffusing capacity for oxygen and carbon dioxide for five classes of tracheae according to their diameters, from 0.2 microm to 35 microm. The harmonic mean thickness of the tracheal epithelium is lowest in smallest tracheae and increases with increasing tracheal diameter. Although the smallest tracheae make up 70% (O2) and 60% (CO2) of the total diffusing capacity, the proximal four classes may also be significant in diffusion of oxygen and particularly of carbon dioxide. The suppression of the development of respiratory pigments in the evolution of terrestrial insects may have increased the relative importance of small tracheal elements for local oxygen consumption.     

Comparative study of the ring glands from wild type and 1(2)gl mutant drosophila melanogaster

Structural and functional changes have been correlated during metamorphic degeneration of a single muscle fiber, the plantar retractor of G. mellonella, its axon, and their junctions to determine which features persist as long as muscle contractility. Changes commence simultaneously in muscle and nerve near cuticular attachments, and spread towards the center. Alterations associated with the muscle, including appearance of collapsed tracheoles and mitochondria with dense bodies, begin late in the last larval instar. Within 12 hours after pupal ecdysis some tracheolar withdrawal occurs, sarcoplasmic reticulum becomes reduced, and many mitochondria have dense bodies, dense membranes, or are enlarged. By 17–19 hours primary myofilaments and striations begin to disappear, microtubules and autophagic vacuole-like bodies appear, and phagocytes invade the muscle. It remains partially contractile upon electrically stimulating its nerve, the ventral nerve, until these changes spread throughout the fiber. Neuromuscular junction changes, including appearance of dense mitochondria and isolation bodies, begin late in the last larval instar. Junctions become fewer, and none remain in those muscle areas where tracheoles, sarcoplasmic reticulum, and primary myofilaments have disappeared. Preliminary studies on nerve discharge activity to the muscle suggest that nerve silence occurs at approximately the time when the muscle loses all contractility. In some axons isolation bodies appear and neurotubules are lost, other axons remain unchanged, and new ones develop later in the pupal state. Phagocytes invade the neural lamella and it disappears in the late pupa, but it reappears in the adult. The adult ventral nerve has over three times more axons and a thinner layer of glial cells than the larval nerve.     

Caterpillars have evolved lungs for hemocyte gas exchange

Since insect blood usually lacks oxygen-carrying pigments it has always been assumed that respiratory needs are met by diffusion in the gas-filled lumen of their tracheal systems. Outside air enters the tracheal system through segmentally arranged spiracles, diffuses along tubes of cuticle secreted by tracheal epithelia and then to tissues through tracheoles, thin walled cuticle tubes that penetrate between cells. The only recognized exceptions have been blood cells (hemocytes), which are not tracheated because they float in the hemolymph. In caterpillars, anoxia has an effect on the structure of the hemocytes and causes them to be released from tissues and to accumulate on thin walled tracheal tufts near the 8th (last) pair of abdominal spiracles. Residence in the tufts restores normal structure. Hemocytes also adhere to thin-walled tracheae in the tokus compartment at the tip of the abdomen. The specialized tracheal system of the 8th segment and tokus may therefore be a lung for hemocytes, a novel concept in insect physiology. Thus, although as a rule insect tracheae go to tissues, this work shows that hemocytes go to tracheae.     

Ventilatory Mechanism and Control in Grasshoppers

SYNOPSIS. Grasshoppers exhibit a diversity of ventilatory patternsdepending on activity status. For each pattern, the mechanismand control of gas exchange is analyzed in terms of a two-stepmodel, consisting of tracheolar and trans-spiracular steps inseries. During the intermittent gas exchange that characterizesthe most quiescent grasshoppers, spiracles open and close inresponse to changing carbon dioxide, and trans-spiracular resistancecontrols gas exchange. In resting but alert grasshoppers, abdominalpumping occurs, and gas exchange is controlled equally by tracheolarand trans-spiracular resistances; tracheal oxygen and carbondioxide are regulated by variation in abdominal pumping andspiracular opening. During hopping, abdominal pumping does notoccur, and bulk gas flow is driven by cuticular deformationsassociated with locomotion. Increased cellular oxygen consumptiondepends on use of internal oxygen stores and increased partialpressure gradients. After hopping ceases, abdominal pumpingincreases dramatically and restores tracheal gas composition;however, the rise in abdominal pumping after hopping is notaffected by tracheal gas levels. During flight, bulk flow tothe flight muscles is driven by tidal thoracic auto-ventilation,while the remainder of the body is ventilated by abdominal pumping.During both hopping and flight, the greatest resistances togas transport exist in the tracheolar rather than the trans-spiracularstep.     

The Lower Devonian scorpionWaeringoscorpio and the respiratory nature of its filamentous structures, with the description of a new species from the Westerwald area, Germany

The fossil scorpionWaeringoscorpio hefteri Størmer, 1970 (Arachnida: Scorpiones) from the Lower Devonian of the Rhenish Massif of Germany is redescribed based on both the original type and newly collected material. A second, more tuberculate species from Siegenian strata near Bürdenbach in the Westerwald (also part of the Rhenish Massif) is described asW. westerwaldensis n. sp. Details of the coxo-sternal region — including the lack of an oral tube — and the number of ventral mesosomal plates are discussed.Waeringoscorpio Størmer, 1970 is best known for its possession of externally-projecting ‘gills’. Our new material reveals that these are indeed pair-wise bundles of rigid, branching filaments which originate laterally, quite possibly from those segments of the mesosoma associated with the book lungs in extant scorpions. Their gross morphology is most consistent with a respiratory organ adapted for use in water. Indeed their closest modern analogues are the tracheal gills of secondarily aquatic insects. We suggest that the morphology and likely palaeoenvironment ofWaeringoscorpio could indicate an aquatic animal, but we draw attention to the uniqueness of its gill-structures, which may not be part of the scorpion ground-pattern. Thus,Waeringoscorpio was perhaps a secondarily aquatic scorpion adapted for benthic life in oxygen-stressed, freshwater-brackish environments.