On the biology of the bird parasite <Emphasis Type="Italic">Neottiophilum praeustum</Emphasis> (Meigen, 1826) (Diptera,Neottiophilidae)

The first data on blood-sucking ectoparasitic larvae of Neottiophilum praeustum (Meig.) which develop in bird nests are presented in Russia, with the fieldfare Turdus pilaris L. as a host example. Larval development takes not more than 10–12 days but no puparia are formed until late autumn. The larvae of Neottiophilum resemble those of calliphorid flies both in body structure and life mode. The main diagnostic characters of Neottiophilum larvae distinguishing them from calliphorid ones are the spiracular disk of the posterior spiracles being positioned dorsal rather than ventral to the stigmal plate and lying outside rather than inside its peritreme. In addition, the anterior spiracles have 14–15, rather than 3–8 spiracular chambers.     

Morphology of spiracles in adult Hyalomma truncatum ticks (Acari; Ixodidae)

Scanning electron microscopical investigations of fractures and corrosion casts of spirales in adult ticks of Hyalomma truncatum revealed a three-part structure consisting of the spiracular plate forming the outer part followed by the subostial space, which leads into the atrial chamber from which the main tracheal trunks arise. The spiracular plate sonsists of a thin surface plate perforated by aeropyles, an underlying interpedicellar space formed by pedicels and an inner thick base plate. The surface plate is subdivided into a porous and a non-porous area. The macula is surrounded by the porous area and cleft by the ostium, which is bounded by a lip. The lip rests on a stalk which passes through the subostial space and forms the lateral wall of the atrial chamber. The interpedicellar space is chambered comprising four types of chambers. Large pyriform chambers (type 1) open to the atmosphere via a large aeropyle and are connected at their base with a duct traversing the base plate. They correspond numerically and in their position with the large aeropyles and the ducts of the base plate. Each chamber is surrounded by four to six medium-sized tubular chambers (type 2) which are closed at both ends. Small tubular chambers (type 3) open to the atmosphere via a small aeropyle, are closed at their base and correspond in number and position to the small aeropyles. Elongated chambers (type 4) are arranged in two to three rows around the subostial space and are closed at both ends. The front row communicates with the subostial space via large gaps. All chambers interconnect with each other by slit-like fenestrations. Below the macula and surrounding the stalk is the subostial space. Over the medial half, the subostial space opens into the atrial chamber. The lateral wall of the atrial chamber is thick, whereas the opposite wall is thin, folded and can be everted and inverted. Inverted, the medial wall closes up the opening to the subostial space and the main tracheal trunks. The base of the atrial chamber sonsists of the openings of the main tracheal trunks only. It is concluded that the aeropyles constitute the functional openings of a spiracle, the interpedicellar space and the subostial space act as diffusion barrier and the atrial chamber is exclusively responsible for the motory process of in- and expiration and is the only closing device of the spiracle.     

Neural control of homologous behaviour patterns in two blaberid cockroaches

Activity patterns of motoneurones which innervate spiracular muscles in two blaberid cockroaches, Blaberus discoidalis and Gromphadorhina portentosa, have been monitored during two homologous behaviour patterns: respiratory and non-respiratory tracheal ventilation. Based upon the activity of spiracular motoneurones during these two activities, the abdominal spiracles have been divided into three functional groups: vestigial, respiratory and non-respiratory. In Blaberus discoidalis spiracle 3 is vestigial, spiracles 6, 7, 8 and 10 are respiratory, and spiracles 4, 5 and 9 are non-respiratory. In Gromphadorhina portentosa spiracles 3 and 10 are vestigial, spiracle 4 is non-respiratory and spiracles 5–9 are respiratory.Respiratory spiracles in both species are characterized by activity patterns of their motoneurones during respiratory tracheal ventilation: low frequency firing at irregular intervals during the respiratory pause and a higher frequency burst synchronous with the expiratory abdominal compression. Non-respiratory spiracles are characterized by complete inactivity of their opener motoneurones during respiratory tracheal ventilation. These motoneurones are activated by mechanical stimulation in both species, which simultaneously suppresses activity in respiratory opener motoneurones. In Blaberus discoidalis, there are no differences between activity patterns of respiratory and non-respiratory closer motoneurones. In Gromphadorhina portentosa, not only do respiratory and non-respiratory closer motoneurones have different activity patterns, but the activity pattern of respiratory closer motoneurones is different during respiratory and non-respiratory tracheal ventilation. The functional implications of these several spiracular motoneurone activity patterns are discussed.     

SPIRACLE STRUCTURE IN TICKS (IXODIDA: ANACTINOTRICHIDA: ARACHNIDA): RÉSUMÉ, TAXONOMIC AND FUNCTIONAL SIGNIFICANCE

Spiracle and tracheal structure in the extant Ixodida is revised and shown to comprise 29 distinct component characters, some of which are common to all Anactinotrichida, while others are unique to the Ixodida or one of its six component clades of ((Argasidae Nuttalliellidae) (Prostriata Metastriata)). Structural variation both between and within families is based upon combinations of minor differences in the component characters, only one of which, spiracular position, proved to be incongruent within the most parsimonious cluster- and tree-analysis solutions. Tracheal airflow in ticks is mediated via passive diffusion gradients. In the argasid spiracle, both aeropyles and ostium are functional, although the latter is only opened briefly during infrequent periods of activity. The ixodid ostium is sealed and all gas exchange takes place via an enlarged sieveplate which reduces transpiration via small aeropyles, an underlying dense array of pedicels and possibly hygroscopic sub-atrial glands. Changes in spiracular morphology from a more ‘ancestral’ argasid type to a more ‘derived’ ixodid type are correlated with changes in tick behaviour, particularly with increased activity associated with the change from nidicoly to host-seeking.     

Respiratory significance of the external morphology of adults and fifth instar nymphs of Notonecta undulata Say (Aquatic Heteroptera; Notonectidae)

The bodies of adult and fifth instar Notonecta possess external air stores which are periodically renewed at the surface of the water. Both nymphs and adults have large ventral air stores on the thorax and abdomen and obtain atmospheric air at the posterior end of the latter; the adult also has dorsal subalar and supra-alar air stores on both these regions. Ten pairs of spiracles open onto the air stores. Although the seven small, ventrally placed abdominal spiracles are probably both exhalant and inhalant in nymphs and adults, the three large anterior spiracles (mesothoracic, metathoracic, and first abdominal), which play a more important respiratory role, appear to function differently in mature and immature Notonecta. In the nymph they are probably both inhalant and exhalant, and communicate broadly with each other and with the ventral air stores. In the adult, however, they open onto separate, air-filled chambers, each of which communicates differently with various parts of the air stores. Although all three probably function in exhalation, only the first abdominal spiracle, whose spiracular chamber is widely continuous with the dorsal and ventral air stores, appears to be well suited for inhalation. Several morphological features, most notably the development of long prothoracic lobes, separate spiracular chambers, and long, movable forewings, allow the adult a greater variety of respiratory modes than are available to the nymph. Some of the respiratory advantages of the adult are: (1) a larger amount of stored air; (2) a longer subalar air store, which can serve as an alternate pathway between the air stores and the atmosphere; (3) a greater capacity to utilize dissolved as well as atmospheric oxygen; (4) greater separation and functional specialization of the three anterior spiracles, thus allowing more separation of exhaled air from oxygen-rich air on the external surface of the thorax; (5) the probable ability to regulate the continuity between various parts of the air stores, thus utilizing alternate pathways of air circulation and/or changing the functions of the three anterior spiracles; and (6) better protection of the latter against the entry of water during prolonged submergence.     

Scanning electron microscopy of the posterior spiracles of cattle grubs Hypoderma bovis and Hypoderma lineatum

Posterior spiracles of newly hatched first instar larvae of Hypoderma bovis (L.) and H. lineatum (DeVill.) consist of two pairs of spiracular openings. Each pair is surrounded by a rima bearing three spines. Posterior spiracles of second instar larvae are composed of a pair of medial ecdysial scars bounded laterally by spiracular plates. H. bovis spiracular plates have twenty-nine to forty openings, each surrounded by a slightly raised rima. H. lineatum spiracular plates have eighteen to twenty-five openings. Spiracular openings lead to posterior felt chambers which are connected to a common anterior felt chamber filled with a meshlike network. In third instar H. bovis each medial ecdysial scar is surrounded by a strongly concave spiracular plate. Spiracular openings are surrounded by slightly raised rima. Most rimae bear a spine. Spiracular plates of H. lineatum are flat and rimae are without spines. Each spiracular opening leads to a posterior felt chamber, several of which are confluent with a larger anterior felt chamber. Anterior felt chambers open into the dorsal longitudinal tracheal trunk. Felt chambers in third instar larvae are also filled with a complex mesh.     

Tradeoffs between metabolic rate and spiracular conductance in discontinuous gas exchange of Samia cynthia (Lepidoptera, Saturniidae)

The insect tracheal system is a unique respiratory system, designed for maximum oxygen delivery at high metabolic demands, e.g. during activity and at high ambient temperatures. Therefore, large safety margins are required for tracheal and spiracular conductance. Spiracles are the entry to the tracheal system and play an important role in controlling discontinuous gas exchange (DGC) between tracheal system and atmosphere in moth pupae. We investigated the effect of modulated metabolic rate (by changing ambient temperature) and modulated spiracular conductance (by blocking all except one spiracles) on gas exchange patterns in Samia pupae. Both, spiracle blocking and metabolic rates, affected respiratory behavior in Samia cynthia pupae. While animals showed discontinuous gas exchange cycles at lower temperatures with unblocked spiracles, the respiratory patterns were cyclic at higher temperatures, with partly blocked spiracles or a combination of these two factors. The threshold for the transition from a discontinuous (DGC) to a cyclic gas exchange (^cycGE) was significantly higher in animals with unblocked spiracles (18.7 nmol g⁻¹ min⁻¹ vs. 7.9 nmol g⁻¹ min⁻¹). These findings indicate an important influence of spiracle conductance on the DGC, which may occur mostly in insects showing high spiracular conductances and low metabolic rates.     

Btk-dependent epithelial cell rearrangements contribute to the invagination of nearby tubular structures in the posterior spiracles of Drosophila

The Drosophila respiratory system consists of two connected organs, the tracheae and the spiracles. Together they ensure the efficient delivery of air-borne oxygen to all tissues. The posterior spiracles consist internally of the spiracular chamber, an invaginated tube with filtering properties that connects the main tracheal branch to the environment, and externally of the stigmatophore, an extensible epidermal structure that covers the spiracular chamber. The primordia of both components are first specified in the plane of the epidermis and subsequently the spiracular chamber is internalized through the process of invagination accompanied by apical cell constriction. It has become clear that invagination processes do not always or only rely on apical constriction. We show here that in mutants for the src-like kinase Btk29A spiracle cells constrict apically but do not complete invagination, giving rise to shorter spiracular chambers. This defect can be rescued by using different GAL4 drivers to express Btk29A throughout the ectoderm, in cells of posterior segments only, or in the stigmatophore pointing to a non cell-autonomous role for Btk29A. Our analysis suggests that complete invagination of the spiracular chamber requires Btk29A-dependent planar cell rearrangements of adjacent non-invaginating cells of the stigmatophore. These results highlight the complex physical interactions that take place among organ components during morphogenesis, which contribute to their final form and function.     

The length of discontinuous gas exchange cycles in lepidopteran pupae may serve as a mechanism for natural selection

Gas exchange is studied in diapausing pupae of Mamestra brassicae L., whose larvae are reared under identical conditions. The release of CO₂ gas is recorded with infrared gaseous analyzers. Oxygen convective uptake into the tracheae and oxygen consumption rates are recorded by means of a constant‐volume coulometric respirometer. Outputs from both of these respirometry systems are combined with infrared actographs. All 3‐month‐old pupae of M. brassicae display a pattern of discontinuous gas exchange (DGE) cycles of CO₂ gas release by bursts, although the lengths of these cycles varies between individuals. Some pupae exhibit long DGE cycles of at least 20 h in duration, with negligible CO₂ gas release during interburst periods, and there is presumed to be a convective gas exchange at this time. As a result of a partial vacuum inside the tracheae, a large oxygen convective uptake always occurs at the start of the spiracular opening phase. Other pupae have short DGE cycles of less than 3 h in duration, with elevated CO₂ gas release during the interburst period, when gas exchange is predominantly diffusive. The spiracular open phase in these pupae consists of frequent separate convective bursts of CO₂ gas release, with the opening–closing rhythms of the spiracles, which are considered as O phase fluttering. The pupae with long DGE cycles exhibit extremely low metabolic rates and very low total water loss rates, whereas those with short DGE cycles have higher metabolic and total water loss rates. The pupae with long DGE cycles live approximately twice as long as those with short cycles; thus, the present study demonstrates that long DGE cycles confer a fitness benefit on pupae as a result of a lower metabolic rate associated with water economy, conferring on them a longer life.     

Spiracle activity in moth pupae—The role of oxygen and carbon dioxide revisited

After decades of intensive research, the actual mechanism behind discontinuous gas exchange in insects has not been fully understood. One open question concerns the actual way (closed, flutter, and open) of how spiracles respond to tracheal gas concentrations. As the results of a classic paper [Burkett, B.N., Schneiderman, H.A., 1974. Roles of oxygen and carbon dioxide in the control of spiracular function in cecropia pupae. Biological Bulletin 147, 274-293] allow ambiguous interpretation, we thus reexamined the behavior of the spiracles in response to fixed, controlled endotracheal gas concentrations.The tracheal system of diapausing pupae of Attacus atlas (Saturniidae, Lepidoptera) was flushed with gas mixtures varying in PO₂ and PCO₂ while the behavior of the spiracles was monitored using changes in the pressure signal. This novel pressure based technique proved to be superior to classic visual observation of single spiracles. A two-dimensional map of the spiracle behavior in response to endotracheal PO₂ and PCO₂ was established. Typically, it contained two distinct regions only, corresponding to “closed” and “open” spiracles. A separate “flutter” region was missing. Because fluttering is commonly observed in moth pupae, we suggest that the intermittent spiracle opening during a flutter phase is an effect of non-steady-state conditions within the tracheal system. For low PCO₂ the minimum PO₂ resulting in open spiracles was linearly dependent upon PCO₂. Above a threshold of 1-1.5 kPa CO₂ the spiracles were open irrespective of PO₂. We propose a hypothetical spiracular control model, which is simple and explains the time course of endotracheal partial pressures during all phases of discontinuous gas exchange.     

Spiracular closing mechanisms in the firebrat, Thermobia domestica (packard) (Thysanura)

Mechanisms for regulating the degree of opening of its spiracles are present in Thermobia. That of the mesothoracic spiracle is of the external type with a flap-like hood guarding the spiracular aperture. Contraction of muscles open the spiracle by raising the hood. Closure is brought about by muscular relaxation and elastic cuticular recoil. Opening is either partial, with small-scale oscillatory movements ('fluttering'), or complete ('wide-opening'). Wide-opening follows bouts of muscular activity. Carbon dioxide anaesthesia relaxes the opener muscles causing the spiracles to close by elastic recoil. This explains continued low tracheal water loss during anaesthesia, and also in death. The control mechanisms of the metathoracic and 8 pairs of abdominal spiracles are of the internal type, with a crypt-like atrium leading into the slit-like neck region of the spiracular pit, one side of which has an elastic cuticular rod running along it. Muscles inserted on the opposite side widen the aperture. As with the mesothoracic spiracle, closure is brought about by muscular relaxation and elastic cuticular recoil.     

Discontinuous gas exchange in dung beetles: patterns and ecological implications

This study correlates a distinctive pattern of external gas exchange, referred to as the discontinuous gas exchange cycle (DGC), observed in the laboratory, with habitat associations of five species of telecoprid dung beetles. The beetles were chosen from a variety of habitats that would be expected to present different amounts of water stress. All five species exhibited DGC. Sisyphus fasciculatus has been recorded only in woodland areas, and does not have strict spiracular control during its DGC. Anachalcos convexus and Scarabaeus rusticus are associated with open mesic habitats. Both species exhibit a distinct DGC, previously found in some other insect species, but intermediate within this study group. Sc. flavicornis and Circellium bacchus are typically found in arid regions, and have the most unusual form of DGC, with spiracular fluttering during the burst phase. These results support the hypothesis that spiracular fluttering reduces respiratory water loss. From this study we conclude that the DGC is an ancestral adaptation, most probably as a result of anoxic environments in underground burrows, but that spiracular control is enhanced to reduce respiratory water loss in beetle species that live in arid habitats. Received 4 August 1999 / Accepted: 7 October 1999     

Comparative micromorphology of third instar larvae and the breeding biology of some Afrotropical Sarcophaga (Diptera: Sarcophagidae)

Four sympatric species of Sarcophaga, viz S. cruentata Meigen, S. exuberans Pandelle, S. nodosa Engel and S. tibialis Macquart, which occur in the Transvaal, South Africa, showed oviparity under optimum laboratory breeding conditions. Details of the life cycle duration under these conditions are discussed. Rearing and colonizing methods were developed. Scanning electron microscopy of third instar larvae provided useful data in distinguishing between the four species. The characters which were examined were the spinulation of the body segments and the rim surrounding the spiracular atrium of the posterior spiracles, the anterior spiracles and the spiracular hairs of the posterior spiracles.     

Morphological aspects of the third instar larva of Haematobia irritans

The main morphological features of the cephalic region of the larva of Haematobia irritans (L.) are the oral grooves, tripartite labium and the antennomaxillary protuberances that have the dorsal, terminal and ventral sensory organs. The total number of sensilla that are found on the terminal organ differs from other cyclorrhaphous-fly larvae. The fan-shaped anterior spiracles usually consist of seven bulbous digits that are unequal in length. The creeping welts consist of notched, convex plates that split into two separate plates as they approach the midline of the venter. This characteristic has not been described previously for this species or other, higher, dipterous larvae. There are two posterior spiracles with an ecdysial scar, four fan-shaped and branching spiracular hairs and irregularly-shaped spiracular openings. The longitudinal anal opening is situated in the cuticular band that is known as the anal organ.     

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。     

Respiratory system of the primitive moth Micropterix calthella (Linnaeus) (Lepidoptera : Micropterigidae)

The 1st thoracic spiracular atrium is closed by anterior and posterior muscle fibres extending between its dorsal and ventral wall. The 2nd thoracic spiracle has only a single (anterior) closing lip, movable by a muscle inserting on the wall below the spiracular aperture; this configuration may be a lepidopteran ground-plan autapomorphy. There are functional spiracles on abdominal segments I – VII, each with a closing “bow” and “lever”. There are intrinsic occlusor muscles in all abdominal spiracles and the 1st spiracle has an extrinsic (ventral) dilator. Dorsal dilator muscles or ligaments are absent. A dorsal and a ventral tracheal trunk extend from the 1st thoracic spiracle into the head; the latter supplies the mouthparts and the antenna; there is no connection between the dorsal and ventral cephalic trunk systems. There is a single series of lateral connectives between the spiracles of each side. There is a ventral tracheal commissure in both pterothoracic segments, but none in the prothorax. In each pterothoracic segment an anterior and a posterior tracheal arch give off branches to the wing and anastomose with each other on their downwards course into the leg. Wing tracheation is greatly reduced. The anterior and posterior tracheae of each wing are independent of each other. There is a dorsal commissure in abdominal segment VIII; ventral abdominal commissures are lacking in Micropterix, although present in other micropterigid genera. The terminalia are partly supplied from tracheae arising in segment VII. Air sacs occur in the tibiae only. Phylogenetic aspects of holometabolan tracheation patterns are discussed.     

Respiratory significance of the thoracic and abdominal morphology of Belostoma and Ranatra (insecta,heteroptera)

Summary Both Belostoma and Ranatra possess I–II, subepimeral, thoracic subalar, and abdominal subalar air stores. In Belostoma, unlike Ranatra, the subepimeral air store is greatly enlarged, the abdominal subalar store is partially exposed to the water, and a fully exposed ventral abdominal air store is also present. All the air stores of Ranatra are normally concealed.The mesothoracic and metathoracic spiracles, which open onto the I–II and subepimeral air stores respectively, are of limited permeability. They appear to have less respiratory importance than the large and highly permeable first abdominal spiracles, which lie in the subalar air space and can probably exhale and inhale large amounts of air. The large eighth abdominal spiracles, which lie at the base of the siphon or retractile organ, can also inhale or exhale much air in Ranatra but appear to be mainly exhalant in Belostoma. The smaller second through seventh abdominal spiracles structurally resemble the eighth ones in Belostoma and open onto the ventral abdominal air store. In Ranatra they appear to have no significant respiratory function.Both genera obtain atmospheric air and give off exhaled air by means of the posterior retractile organ or siphon. The two types of air appear to follow different pathways in the two genera. In Ranatra atmospheric air appears to enter the tracheal system mainly or entirely through the eighth abdominal spiracles and then passes through the first abdominal spiracles into the subalar space. Exhaled air follows the reverse pathway. In Belostoma, however, atmospheric air probably enters the tracheae mainly through the first abdominal spiracles; it is conveyed to these spiracles from the retractile organ through the subalar space or, more indirectly, through the ventral abdominal air store. Air exhaled through the first abdominal spiracles follows the reverse route; the eighth abdominal spiracles can also exhale directly into the base of the retractile organ.During underwater respiration the abdominal portion of the subalar air store appears to be the main reservoir for oxygen. The subalar oxygen is initially atmospheric, and is supplemented, during submersion, by other sources of oxygen. Belostoma may use its exposed ventral abdominal air store, and its partially exposed abdominal subalar one, as physical gills; both these stores communicate with the inhalant first abdominal spiracles. Ranatra, none of whose air stores are normally exposed, appears, to be less capable of utilizing dissolved oxygen, but the considerable amount of atmospheric oxygen in the elongated siphon may be inhaled, during submersion, through the eighth abdominal spiracles.In both genera the thoracic air stores appear to be of less respiratory importance than the abdominal ones. They do not appear capable of obtaining large amounts of oxygen, and the thoracic spiracles are relatively impermeable. All the air stores, however, serve to protect the spiracles against the entry of water, and also contribute to the body's hydrostatic balance. It is also possible that some of the air stores play a role in pressure reception.The literature indicates much intergeneric variation in the respiration of Belostomatidae and Nepidae. In the Belostomatidae there is considerable variation in the extent of the ventral abdominal air store and in the roles of the subalar air store and the spiracles. The Nepidae show differences in their ability to utilize dissolved oxygen and in the extent of the subepimeral air store.