Larval morphology of the lesser housefly, Fannia canicularis

The morphology of all larval instars of Fannia canicularis (Linnaeus) (Diptera: Fanniidae) is documented using a combination of light and scanning electron microscopy. The following structures are documented for all instars: antennal complex; maxillary palpus; facial mask; cephaloskeleton; ventral organ; anterior spiracle; Keilin's organ; posterior spiracle; fleshy processes, and anal pad. Structures reported for the first time for all instars include: two pairs of lateral prominences on the prothoracic segment; additional ventrolateral prominences on the second thoracic segment, and a papilla at the base of the posterior spiracle. Other structures reported for the first time are anterior spiracles in the first instar and a serrated tip on the mouthhook in the second instar. A trichoid sensillum on the posterior spiracular plate, representing a sensory organ otherwise unknown in the Calyptratae, is described in the second and third instars. Results are discussed and compared with existing knowledge on dipteran larval morphology.     

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.     

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.     

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.     

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.     

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。     

普通齿蛉幼虫的呼吸系统及呼吸行为

研究了普通齿蛉Neoneuromus ignobilis Navás幼虫的呼吸系统及其呼吸行为。结果表明:普通齿蛉幼虫为全气门式(10对气门)呼吸系统,前中胸、中后胸之间、腹部8节各有1对气门,腹部8节各有气管鳃1对,前6对细短,管状,有较短绒毛,后2对气管鳃较粗长,呈羽毛状。腹部1～7节各有1对毛簇,第8腹节无毛簇。侧纵干气管较粗,4束,自前胸前缘部分成左右2组,每组两根侧纵干气管,向胸腹部延伸,二级气管分别伸达各个气门和毛簇,腹部每节由毛簇处的二级气管分支而来的三级气管相连或延伸至消化道等处。气管鳃中无气管。有毛簇呼吸、气门呼吸和体壁呼吸3种呼吸方式,在水中以毛簇呼吸为主,在陆上进行气门呼吸和体壁呼吸。     

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.     

Morphology of the first instar of Calliphora vicina, Phormia regina and Lucilia illustris (Diptera, Calliphoridae)

Scanning electron microscopy documentation of first instar Calliphora vicina Robineau-Desvoidy, Phormia regina (Meigen) and Lucilia illustris (Meigen) (Diptera: Calliphoridae) is presented for the first time, and the following morphological structures are documented: pseudocephalon; antenna; maxillary palpus; facial mask; labial lobe; thoracic and abdominal spinulation; spiracular field; posterior spiracles, and anal pad. Light microscopy documentation and illustrations are provided for the cephaloskeleton in lateral and ventral views. New diagnostic features are revealed in the configuration of the facial mask, cephaloskeleton and posterior spiracles. The first instar morphology of C. vicina, Ph. regina and L. illustris is discussed in the light of existing knowledge about early instars of blowflies.     

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.     

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.     

Expression of a surface epitope on cells that link branches in the tracheal network of Manduca sexta.

A monoclonal antibody (MAb 2F5) to a cell surface epitope labels a small subpopulation of tracheal epithelial cells in each thoracic and abdominal segment of Manduca. These cells (nodes) represent the sites within the tracheal network at which invaginating tracheal tubes join during embryonic establishment of the tracheal network. Tracheal nodes are also the sites at which tracheal cuticle fractures during each molt. Since tracheal cuticle is shed through each spiracle, a tracheal node lies between each pair of contralateral spiracles within a segment (commissural node) and between each pair of adjacent, ipsilateral spiracles (lateral longitudinal node). MAb 2F5 first labels presumptive nodal cells of tracheal epithelium immediately prior to the linking of epithelial tubes from successive and opposite spiracles. One cell at the tip of each invaginating tracheal branch labels with MAb 2F5. The highly localized expression of the cell surface epitope recognized by MAb 2F5 may be instrumental in the orderly coupling of tracheal branches during embryonic development. On the basis of immunolabeling of Western blots and tissues, MAb 2F5 is believed to recognize Manduca fasciclin II, a member of a class of molecules involved in cell adhesion/recognition.     

Regulation and metamorphosis of the abdominal histoblasts ofDrosophila melanogaster

Summary The development of the adult abdomen ofDrosophila melanogaster was analyzed by histology, microcautery, and genetic strategies. Eight nests of diploid histoblasts were identified in the newly hatched larva among the polytene epidermal cells of each abdominal segment: pairs of anterior dorsal, posterior dorsal, and ventral histoblast nests and a pair of spiracular anlagen. The histoblasts do not divide during larval life but begin dividing rapidly 3 h after pupariation, doubling every 3.6 h. Initially they remain confined to their original area, but 15 h after pupariation the nests enlarge, and histoblasts replace adjacent epidermis cell by cell. The histoblasts cover half the abdomen by 28 h after pupariation and the rest by 36 h. Polytene epidermal cells of the intersegmental margin are replaced last. Cautery of the anterior dorsal nest caused deletion of the whole corresponding hemitergite, whereas cautery of the posterior dorsal nest caused the deletion of the macrochaetae of the posterior of the hemitergite. Cautery of the ventral nest deleted the hemisternite and the pleura, whereas cautery of the spiracular anlagen deleted the spiracle. Results of cautery also revealed that no macrochaetae formed on the tergite in the absence of adjacent microchaetae. Clonal analysis revealed that there were no clonal restrictions within a hemitergite at pupariation. Cautery of polytene epidermal cells other than those of the intersegmental margin failed to affect tergite development. However, cautery of polytene epidermal cells of the intersegmental margin adjacent to either dorsal histoblast nest caused mirror-image duplications of the anterior or posterior of the hemitergite in 10% of the hemitergites. Forty percent of the damaged presumptive hemitergites formed complete hemitergites, indicating extensive pattern regulation and regeneration. Pattern duplication and regeneration were accounted for in terms of intercalation and a model of epimorphic pattern regulation (French et al., 1976). Histoblasts in adjacent segments normally develop independently, but if they are enabled to interact by deleting the polytene epidermal cells of the intersegmental margin, they undergo intercalation which results in duplication or regeneration. The possible role of the intersegmental margin cells of insects in development was analyzed.     

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.     

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.     

Tracheal ultrastructure of protura (Insecta : Apterygota)

The tracheal systems of Sinentomon and Eosentomon (Apterygota : Protura) were examined in thin sections and compared with the tracheae of collembolan, Allacma. The tracheal system of Protura consists of spiracles and tracheae. The spiracle is a simple, concave cuticular cavity known as an atrium. A globular chamber is present between the atrium and trachea. The atrium of Eosentomon is decorated with ridges and has 2 small openings to tracheal recesses beside the central tracheal opening. The tracheae of Protura are characterized by a high frequency of taenidia and the absence of intima folds and intertaenidial spaces. The taenidia of Sinentomon have a rectangular section and those of Eosentomon are gable-shaped. The results also suggest that the tracheal recess of Eosentomon is a kind of stigmatic gland. The tracheal structure of Protura was compared with that of collembolans, insects, and other arthropods, and discussed in terms of phylogeny.     

SALMFamide2 and serotonin immunoreactivity in the nervous system of some acoels (Xenacoelomorpha)

Acoel worms are simple, often microscopic animals with direct development, a multiciliated epidermis, a statocyst, and a digestive parenchyma instead of a gut epithelium. Morphological characters of acoels have been notoriously difficult to interpret due to their relative scarcity. The nervous system is one of the most accessible and widely used comparative features in acoels, which have a so‐called commissural brain without capsule and several major longitudinal neurite bundles. Here, we use the selective binding properties of a neuropeptide antibody raised in echinoderms (SALMFamide2, or S2), and a commercial antibody against serotonin (5‐HT) to provide additional characters of the acoel nervous system. We have prepared whole‐mount immunofluorescent stainings of three acoel species: Symsagittifera psammophila (Convolutidae), Aphanostoma pisae, and the model acoel Isodiametra pulchra (both Isodiametridae). The commissural brain of all three acoels is delimited anteriorly by the ventral anterior commissure, and posteriorly by the dorsal posterior commissure. The dorsal anterior commissure is situated between the ventral anterior commissure and the dorsal posterior commissure, while the statocyst lies between dorsal anterior and dorsal posterior commissure. S2 and serotonin do not co‐localise, and they follow similar patterns to each other within an animal. In particular, S2, but not 5‐HT, stains a prominent commissure posterior to the main (dorsal) posterior commissure. We have for the first time observed a closed posterior loop of the main neurite bundles in S. psammophila for both the amidergic and the serotonergic nervous system. In I. pulchra, the lateral neurite bundles also form a posterior loop in our serotonergic nervous system stainings.     

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.     

Scanning electron microscopy of Drosophila melanogaster embryogenesis: III. Formation of the head and caudal segments

The formation of both the anterior most and posterior most segments in higher dipteran embryos involves complex movements of primordia which can be best visualized with the scanning electron microscope. During head formation, the gnathocephalic segments partially involute through the stomodeum. The labial segment forms the floor of the mouth, and the fused maxillary and mandibular segments form the lateral sides of the mouth. The involuted clypeolabrum forms the roof of the mouth. Invaginations of cells for segmentally derived sense organs can be found prior to involution on all the gnathocephalic and thoracic segments as well as on the labrum. The antennal sense organ derives from the lateral surface of the procephalic lobe. Following involution of the mouth parts, the dorsal ridge, which arises just anterior to the first thoracic segment, is drawn over the dorsal procephalic lobe producing the deep dorsal sac. The optic lobes of the brain invaginate anterior to the dorsal ridge just prior to the covering over of the head. The formation of the anal segment is similarly complex. Two rudimentary segments are found posterior to the eighth abdominal segment. During shortening of the germ band, the posterior most segment is drawn around the posterior tip of the embryo to lie ventrally. Two large anal pads form lateral to the anus from this segment. The next segment, following dorsal closure, produces a pair of anal sense organs and a central tuft of setae. Finally, the eighth abdominal segment gives rise to the posterior spiracles. Following dorsal closure these three segments fuse to produce the terminal (anal) segment of the larva.